British Ornithologists’ Club
THE NATURAL
HISTORY MUSEUM
1 2 DEC 2012
PRESENTED
TRING LIBRARY
Volume 132 No. 4
December 2012
FORTHCOMING MEETINGS
See also BOC website: http://www.boc-online.org
BOC MEETINGS are open to all, not just BOC members, and are free.
Evening meetings are held in an upstairs room at The Barley Mow, Horseferry Road, Westminster, London
SW1P 2EE. The nearest Tube stations are Victoria and St. James's Park; and the 507 bus, which runs from
Victoria to Waterloo, stops nearby. For maps, see http://www.markettaverns.co.uk/the__barley_mow.html or
ask the Chairman for directions.
The cash bar will open at 6.00 pm and those who wish to eat after the meeting can place an order. The talk
will start^at 6.30 pm and, with questions, will last about one hour.
It would be very helpful if those who are intending to come would notify the Chairman no later than the day before the
meeting and preferably earlier.
26 February 2013 at 6.30 pm— Dr James Reynolds— Ascension Island and Sooty Terns: an ecological disaster
or a smorgasbord in the eyes of conservation biologists?
Abstract: Ascension Island, a remote 9,700 ha-island in the South Atlantic, is a very important breeding site for
many seabird species. Sooty Tern Onychoprion fuscatus is globally of Least Concern, but its circum-equatorial
distribution means that breeding sites such as Ascension accommodate very large colonies. Subject to severe
predation from introduced species such as Domestic Cats Felis silvestris catus and Common Mynas Acridotheres
tristis, the eradication of the former was expected to result in population recovery. However, numbers remain
stubbornly deflated (i.e. only some 350,000 birds!) compared with a population of two million birds only some
50 years ago. I will explore the pressures on the current population, describe conservation efforts over the last
20 years and present some new findings from tracking of terns that might help us to restore the island to its
former seabird glory.
Biography: Jim Reynolds undertook a D.Phil. at the Edward Grey Institute, University of Oxford, on Uptake and
disposition of calcium and water by egg-laying Zebra Finches. After a period as a Post-doctoral Research Fellow at
the University of Memphis & Archbold Biological Station, USA, for the past ten years he has been a lecturer
in Ornithology and Animal Conservation at the University of Birmingham's Centre for Ornithology, where
he runs their M.Sc. course.
21 May 2013—5.45 pm Annual General Meeting followed at 6.30 pm by several short talks
If you wish to give a talk, which should last 10-15 minutes, please send details to Robert Prys-Jones (e-mail:
r.prys-jones@nhm.ac.uk) no later than 14 January 2013.
Details of the talks on 24 September 2013 and 19 November 2013 will appear in the March Bulletin
Saturday 6 April 2013— Joint meeting with the African Bird Club and the Natural History Museum
A one-day meeting in the Flett Theatre, Natural History Museum, South Kensington, London SW7 5BD.
The provisional programme includes:
• Saving Ethiopia's most threatened endemic bird in a constantly changing environment — Bruktawit Abdu
• Libya's Lesser Crested Terns: a vulnerable and important population — Abdulmaula Hamza
• Birds and birdwatching in Rwanda: a gem in the heart of Africa — Jason Anderson
• Speciation in African white-eyes — Siobhan Cox
• In Archer's footsteps: birding in the Republic of Somaliland — Nigel Redman
• Reconstructing the lost world of the dodo; the extinct birds of the Mascarenes— Julian Hume
The Chairman: Helen Baker, 60 Townfield, Rickmansworth, Herts. WD3 TDD, UK. Tel. +44 (0)1923 772441
E-mail: helen.baker60@tiscali.co.uk
Club Announcements
225
Bulletin of the
BRITISH ORNITHOLbUTST^-dLUB
THE NATURAL
history musI0|*ib O-C
1 2 DEC 2012
PRESENTED
TRING LIBF
2012 132(4)
Vol. 132 No. 4
Published 1 December 2012
CLUB ANNOUNCEMENTS
Chairman's message
Next meeting— Tuesday 26 February 2013 at the Barley Mow, Westminster
The meeting on 18 September 2012 was our last at Imperial College. Next year, starting on Tuesday 26
February, Club meetings will be held in the upstairs room at the Barley Mow, Horseferry Road, Westminster,
London SW1P 2EE. Details are given opposite. UK members should please note that in future there will be no
flyer for booking supper. Those who want to eat after the talk can place an order when they arrive. Also, it
will not be necessary to book in advance, though to give us an idea of numbers and enable us to have name
badges ready, it would be very helpful if those who are intending to come would let me know no later than
the day before the talk and preferably earlier.
The Biodiversity Heritage Library is making good progress adding the Bulletins to its website at www.
biodiversitylibrary.org/bibliography/46639. Most are now freely available on the site. For the latest position
please check the Club news page on the BOC website.
We have recently made a number of changes to our website in an effort to keep it up to date and informative.
Please take a look at it if you have not done so recently.
Helen Baker
REFEREES
I am grateful to the following, who have reviewed manuscripts submitted to the Bulletin during the last year
(those who refereed more than one manuscript are denoted by an asterisk in parentheses): Ignacio Areta,
Norbert Bahr, Richard C. Banks (*), Walter Boles, Michael Brooke, W. R. P. Bourne, Caio Carlos (*), Peter
Castell, David Christie, William S. Clark (*), Mario Cohn-Haft, Martin Collinson, Jose Luis Copete, Pierre-
Andre Crochet, Jon Curson, Geoffrey Davison, Edward C. Dickinson (*), Tulio Dornas, Robert J. Dowsett (*),
Fran^oise Dowsett-Lemaire, Jack Dumbacher, Renate van den Elzen, Rosendo Fraga, Juan F. Freile, Richard
Garrigues, Peter J. Garson, Steve M. Goodman, Floyd Hayes, Julian Hume, Tim Inskipp, Michael C. Jennings,
Leo Joseph, Alan C. Kemp, Alan Knox, Neils Krabbe, Frank Lambert, Daniel F. Lane, Mary LeCroy (*), Huw
Lloyd, Wayne Longmore (*), Phil McGowan, Jose Fernando Pacheco, Eric Pasquet, Richard F. Porter, Doug
Pratt, Robert Prys-Jones (*), Mark B. Robbins, Marcos Rodrigues, Kees Roselaar (*), Eleanor Rowley, Roger
Safford, Cesar Sanchez, Richard Schodde (*), Thomas S. Schulenberg, Frederick Sheldon, Frank Steinheimer
(*), Fernando Costa Straube, Lars Svensson (*), Till Topfer, Colin Trainor, Magnus Ullman, David R. Wells,
Andrew Whittaker (*), Gary Wiles, Jim Wiley, Christopher C. Witt, Iain Woxvold, Kevin J. Zimmer and
Bernard Zonfrillo.— The Hon. Editor
Hadoram Shirihai
226
Bull. B.O.C. 2012 132(4)
Correcting the identification of two rare wheatear records
in Israel
by Hadoram Shirihai
Received 1 December 2011
Summary.— -Basalt Wheatear Oenanthe lugens warriae is one of the least known
of its genus, but recent advances in knowledge of its characters have enabled
records of Variable Wheatear O. picata opistholeuca and Black Wheatear O. leucura
in Israel to be reidentified as O. 1. warriae. Separating the three taxa is discussed,
and the behaviours of Israeli Basalt Wheatears described, providing the first
such data away from the breeding range. Israeli records also suggest that warriae
often undertakes long dispersal, which is supported by the discovery of two
misidentified specimens in Berlin, whose data are presented for the first time here.
Details of a Turkish record of Basalt Wheatear, a bird that hybridised with Finsch's
Wheatear O.finschii are also provided.
The recent description of Basalt Wheatear Oenanthe lugens warriae (endemic to the
basalt deserts of Jordan and Syria; Shirihai et al. 2011) has led to a better understanding
of its morphology, variation and identification. Field separation of O. /. warriae from male
Variable Wheatear O. picata opistholeuca (treated as a morph in some literature), which
mostly breeds in Central Asia, and the geographically well-separated Black Wheatear O.
leucura (of Iberia and north-west Africa) are complex issues. Work on Shirihai et al. (2011)
and related projects (Shirihai & Svensson in prep., Shirihai et al. in prep.) has established
that warriae, opistholeuca and leucura represent a considerable identification challenge.
In Israel, the 1980s and early 1990s were marked by a core of c.15 young birders,
among them Y. Baser, A. Ber, A. Ben Dov, E. Dovrat, Y. Golan, O. Horin, R. Mizrachi, E.
Shochat and myself. From a European perspective, activity mostly centred on the migration
hotspot of Eilat in the south, to which thousands of birders flocked each spring. The Eilat
International Bird watching Centre was founded in 1984, but prior to that the above-
mentioned observers made casual visits to Eilat, mainly between summer and winter,
especially to study migration and to find vagrants.
One 'surprise' was a 'Black Wheatear', found by E. Dovrat in fields at Kibbutz Eilot,
near Eilat, on 11-15 December 1982, and photographed by A. Beer et al. Secondly, a male
'Variable Wheatear' was seen on 4 February 1986 by M. Gellert, H. Fleldbjerg & HS at Eilat
saltpans (Shirihai 1999). It was not photographed and only a written description is available.
However, both were misidentified Basalt Wheatears.
'Black Wheatear'
The following is based on E. Dovrat's field notes (transcribed from Hebrew), two
images by A. Beer (Figs. 1-2) and the brief description in Shirihai (1996).
Size.— Some observers (including myself) considered it larger than the nearby White-
crowned Wheatear O. leucopyga, and even estimated the bird to be as large as Hooded
Wheatear O. monacha. However, Dovrat described it as follows: '[no] real difference in size
compared to leucopyga'. Thus, discrepancy existed among the observers concerning size,
suggesting this was not easy to determine reliably. O. leucura is generally larger and heavier
looking than warriae, being especially larger headed, stronger / longer billed and legged.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
227
Bull. B.O.C. 2012 132(4)
However, warriae is on average c.5% larger in wing and tail than Mourning Wheatear O.
1. lugens, similar to O. leucura. Mean wing length of male O. /. warriae is 98.3 mm, the tail
64.8 mm, compared to male leucura 99.8 mm and 64.3 mm, respectively (Shirihai et al. 2011,
Shirihai & Svensson in prep.).
All of the observers lacked experience with leucura and warriae, and the latter's
characters, including size, were then unknown. We also did not consider the effect of
strong desert light, which often makes birds appear larger, or that in winter, especially in
early morning, birds often 'puff' out their body feathers and appear larger or heavier (at
least temporarily). Thus, identification based on size was both inconstantly judged and
mistakenly afforded too much import.
Structure. — Separating warriae and leucura rests on the former's distinctly longer
primary projection, roughly equal to tertial length (rather than at most c.% of tertial length
in Black). The combination of longer primary projection and on average broader tail-band
in warriae (c.17 mm on outer web of r6, vs. c.14 mm in leucura) is consistently diagnostic.
The wingtip of warriae reaches the tail-band's upper border (at most the gap between them
is as wide as the tail-band itself). In Black they are well separated, with the distance almost
twice that of the tail-band (due to the shorter primary projection and narrower tail-band).
Compare Figs. 1 and 3 with Fig. 5. If the band is invisible, it is still possible to appreciate
Basalt's longer primary extention because the wingtip normally reaches along the
length of the black central rectrices (in Black the wingtip reaches only to where these are
first exposed below the white uppertail-coverts; cf. Fig. 5 vs. Fig. 3). Despite this feature's
utility, caution is needed: it is best judged on perched birds when the wing and tail are held
straight and parallel. The Eilat bird clearly showed these pro-Basalt features (Fig. 1).
These wheatears can also be separated by wing formula, with pi in leucura considerably
longer than the tip of the primary-coverts (in both races pi > 4-11 mm, and usually >6 mm;
pi > 0.0-4. 3 mm or < 0.5 mm in warriae). In flight photographs, the long outermost primary
in Black is often detectable, but not in Basalt. The bird in Fig. 2 shows no indication of a
long pi.
The bill of warriae also averages shorter and thinner, appearing straighter and less
broad-based than in most leucura. Again, the Eilat bird clearly had a shorter / slender bill.
Unique in wheatears, however, the general jizz of Black Wheatear often suggests a 'small
thrush' rather than a wheatear, due to the proportions of the body, head and bill.
Plumage and wing pattern.— Dovrat noted 'In flight a bright [= whitish-grey] tone
across the mid wing... white 'flashes' on underwing brighter than on upperwing and
encompassed a larger area, thus most of underwing looked very bright (the brightest area),
emphasising the clear black shafts; on both wing surfaces the bright panel was surrounded
by black (broadest on the trailing edge), giving a 'frame' effect; Y. Baser, O. Horin, R.
Mizrachi & HS also reported that the white wing pattern recalled Mourning Wheatear.'
This description and the photograph (Fig. 2) best fit a young male warriae (Fig. 4), which
often has relatively less white on the inner webs of the remiges than adult males (Shirihai
et al. 2011). Male leucura often also has dull silvery white undersides to the remiges (Fig. 6),
but this colour is more uniformly distributed over the underwing. The white inner webs
on Basalt are narrow and mostly concealed but sometimes visible in flight. Young Basalt
(especially females) can lack any such white edges or they are mostly concealed, but there is
still a 'wing flash', enhanced by the dark trailing edge and mostly visible from below. Thus,
the description of lugens- like white 'flashes' confirms that the Eilat bird had the diagnostic
Basalt wing pattern.
Overall, the plumage of warriae and leucura are near-identical. However, first-year
Basalt often still has white tips / spots on the primary-coverts (and frequently the alula and
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
228
Bull. B.O.C. 2012 132(4)
tertials) which Black lacks. However, Shirihai et al. (2011) showed that, from late autumn,
adult and some younger Basalt could have much-reduced, or lack, white spots. The Eilat
bird seems to have been one such individual.
Conclusion. — Because warriae and leucura are close in size, this character (which was
inconsistently judged) cannot be used. In all other features the bird was obviously a Basalt
Wheatear. At the time (1982) Basalt Wheatear was effectively unknown, and in effect the
observers had to choose between male Black and Variable Wheatears. Identification as
the former was unsurprising, given that Basalt can be almost as large as Black Wheatear.
Finally, it can safely be separated from male opistholeuca especially by the longer primary
projection but relatively short tail, tail and wing patterns, and wing formula.
'Variable Wheatear'
No photographs exist of this bird, observed at Eilat on 4 February 1986, so the only
material for comparison is my detailed description (summarised in Shirihai 1999).
Size. —Described as 'dearly smaller than White-crowned ... marginally larger and
heavier than Mourning Wheatear', or 'appreciably stockier than Mourning'. O. 1. warriae is
on average rather notably larger than O. 1. lugens : e.g., male Basalt averages c.16 cm, with
wing 98.3 mm and tail 64.8 mm, whereas male O. 1. lugens is c.15.3 cm, 93.1 mm and 60.3
mm, respectively. In overall size (15 cm) and wing (93.1 mm) opistholeuca is closer to O.
1. lugens, but has a proportionately longer tail (67.3 mm) than either (Shirihai et al. 2011,
Shirihai & Svensson in prep.). Thus in size the bird corresponds better with O. 1. warriae,
being larger than O. /. lugens and opistholeuca.
Structure.— My field notes state that overall proportions were much like Mourning and
Finsch's Wheatears O. finschii, with no indication of a proportionately longer tail, which
affords Variable Wheatear a slimmer appearance than 0. 1. lugens or warriae. Thus, structure
/ jizz also favour Basalt Wheatear.
Plumage and wing pattern.— My description reads: 'the remiges appeared completely
dark in flight, save for a grey basal tinge on the underwing . . . the bird was in fresh plumage
and probably an adult, yet there was no trace of the white-tipped primary-coverts normally
visible on adult Basalt Wheatear at this time of year; and the black on the belly was clearly
Legend to figures on page opposite
Figures 1-2. First Israeli record of Basalt Wheatear Oenanthe lugens warriae, Eilat, December 1982 (A. Beer).
Originally misidentified as Black Wheatear O. leucura, note the narrow, pointed bill, long primary projection
(upper arrows in Fig. 1) with wingtip almost reaching upper border of black tail-band (lower arrow),
extensive pale wing 'flashes' enhanced by dark trailing edge, and contrast between pale inner webs and dark
shafts, all of which features are diagnostic of warriae.
Figure 3. Basalt Wheatear Oenanthe lugens warriae, Kfar Barukh, Jezreel Valley, northern Israel, January 2010
(U. Makover). Note similarity to 1982 bird, especially narrow bill, long primary projection (distance between
upper two arrows), broad black tail-band with the upper border of the latter falling level with the wingtip
(rear upper and lower arrows).
Figure 4. Fourth Israeli Basalt Wheatear Oenanthe lugens warriae, 'km 32' north of Eilat, December 2001 (Y.
Perlman)
Figures 5-7. Black Wheatear Oenanthe leucura, Spain (D. Jefferson, top, R. Armada, middle, and, M. Ullman
bottom). Unlike O. lugens warriae, the primary projection (distance between upper two arrows. Fig. 5) is
relatively short, the wingtip just reaching the point where the dark central rectrices are exposed (lower left
arrow). In consequence, the wingtip falls further from the (narrower) black tail-band (distance between the
upper right and lower right arrows). Note the rather strong bill and long pi (Fig. 6). In flight. Black Wheatear
has more uniform and diffuse dull silvery remiges, lacking the more contrasting pattern (especially from
below) of Basalt Wheatear.
Figures 8-14. First-summer, probably female Basalt Wheatear Oenanthe lugens warriae, Ovda Valley, Israel,
March-April 2012 (H. Shirihai)
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
—
Hadoram Shirihai
229
Bull. B.O.C. 2012 132(4)
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
230
Bull. B.O.C. 2012 132(4)
demarcated from the white rear belly (in Variable Wheatear, the black usually extends
further beyond the legs and gradually merges into the white vent, although this character
varies individually and according to posture)/
Given recent advances in our knowledge of warriae and its separation from male
opistholeuca (Shirihai et al. 2011), this description does not eliminate either. Some young
Basalt lose the white tips to the primary-coverts quite early, yet still possess juvenile flight
feathers with no or very narrow white edges to the inner webs, but for a pale grey tone
that is most noticeable on the underwing. This combination best fits the bird in question.
Furthermore, the border between the black and white in the ventral region is now known
to be highly variable, but the fact that the demarcation was well defined favours Basalt.
Contra Shirihai (1999), the demarcation in warriae is better defined and less likely to extend
onto the sides of the undertail-coverts as tends to be true in opistholeuca (Shirihai et al. 2011).
Tail-band. —The description reads: Tts smaller size, broad T-shaped tail-band and
more extensively black belly (reaching just beyond the thighs) eliminated immature White-
crowned'. The tail-band was very broad and I did not notice any indication of uneven black
areas on the webs of the outer rectrices, or that the black on the outer web averaged longer,
like Pied O. pleschanka or Black-eared Wheatears O. hispanica as is often partially true of
opistholeuca. Again, the description better conforms to warriae.
Behaviour.— On landing 'it bobbed up and down— bowing low and then standing
quite upright— while frequently cocking and flicking its tail' (i.e. like Finsch's Wheatear).
This is the only clue that perhaps favours Variable Wheatear, but recent observations
suggest O. 1. lugens and warriae also 'bow' and tail-lift to some extent, but more slowly
and less dramatically than Finsch's, as observed in winter in Israel. Nevertheless, a Basalt
Wheatear in March-April 2012 (Fig. 13) cocked and flicked the tail almost as vigorously as
Finsch's. Retrospectively, I cannot recall that the Eilat wheatear behaved differently from
'extreme' examples of O. /. lugens or warriae.
Conclusion.— Given variation in warriae and its separation from male opistholeuca
(Shirihai et al. 2011), the description could correspond to either taxon, but better fits a
young male Basalt. It lacks any features uniquely suggestive of male opistholeuca. The
misidentification was, again, the result of limited knowledge of warriae at the time. Also,
crucial diagnostics of male opistholeuca (like primary spacing and length of pi vs. the tips
of the primary-coverts) were then unknown. With hindsight I consider this bird to have
been warriae, but in the absence of documentation it is best regarded as a 'presumed Basalt
Wheatear'.
The 1986 bird was definitely not a 'black morph' O.finschii, a possibility also raised with
respect to a Basalt Wheatear in Turkey in May 2011, which was reported to have successfully
bred with a Finsch's Wheatear (see below).
These corrections have been approved by the Israeli Rarity and Distribution Committee,
which has decided to remove Black and Variable Wheatears from the official Israeli list.
Records of Basalt Wheatear in Israel
There have been six records of (and seven individual) Basalt Wheatears in Israel. These
provide the first data concerning dispersal and habitat preferences of warriae away from the
breeding grounds. 11-15 December 1982: one (apparently young, possibly male) just north
of Eilat, in modified gravel desert with cultivated fields nearby; the bird used tall fence posts
as lookouts (Figs. 1-2). 4 February 1986: one (apparently young, possibly male) at Eilat, near
the saltpans, sometimes foraged by the palm planation. December 1994: two in the Arava
Valley (c.80 km north of Eilat) in flat desert adjoining gravel hillsides and fences (cf. Shirihai
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
231
Bull. B.O.C. 2012 132(4)
1996). 6-21 December 2001: first-winter male at 'km 32' north of Eilat (J. P. Smith et al.) on
open sandy plains with some bushes (Fig. 4). 19 January 2010: first-winter male near Kfar
Barukh reservoir, Jezreel Valley (U. Makover; Fig. 3). 21 March-5 April 2012: first-summer,
probably female, Ovda Valley, southern Negev, north of Eilat (D. Berkowic et al.) in a desert
wadi with bushy cover (Figs. 7-13).
December 2001 bird ( per J. P. Smith & Y. Perlman): in patch of Haloxylon bushy habitat
(amid barren desert), selecting notably high perches (c.l m above ground), but also stones
on ground. Generally 'chat-like', using bushes as lookouts to swoop down on ground-strata
prey (insects), spreading the wings and tail on landing, like White-crowned Wheatear. The
bird mostly fed alone. It was silent throughout. Observed in mid afternoon (14.10 h) when
very active. Showed some aggression to a Blackstart Cercomela melanura, and seen once
feeding on fruit of Ochradenus baccatus. Hovered, including several times when collecting
something from a bush.
January 2010 bird (per U. Makover): found on a small side road near Kfar Baruch
reservoir; alone and rather shy, mostly fed on open ground. Silent.
March-April 2012 bird (A. Balaban, I. Berger, A. Ben Dove, B. Granit, D. Berkowic, D.
Forsman, Y. Perlman, I. Shanni, HS et al.): consensus existed among the above-mentioned
observers, and many others familiar with O. 1. lugens, that the bird had a quite different jizz,
foraging and territorial behaviours. Prolonged observations were made on 22 and 26-30
March 2012. Especially on 22nd, the bird constantly bowed (crouching almost 90° forward
and lifting the tail), somewhat like O. finschii (O. 1. lugens may bow but usually less deeply
and constantly). Bowing apparently is a territorial signal used, among other contexts, in
the presence of observers, and appears to decrease with time as the bird adapts to human
presence. To feed it often hovered above ground. Once it mobbed a small snake together
with a Blackstart and Black-eared Wheatear (the warriae detected the snake first, attracted
the other birds and called most persistently, frequently hovering above it, and followed the
snake longest, for c.2 minutes). On 22 March we trapped the bird; on release it gave a short
song sequence (in escape flight), which sounded faster and more musical than lugens. On
all days, especially 30 March, it constantly fed around the bases of flowering Zilla spinosa
bushes, mostly on small insects. Remained 1-3 minutes (once 11 minutes) within a single
bush and in the bushes sometimes c.50% of a given observation period. In 35 years, I have
never seen any wheatear forage in this way. This plant only lives a few years and grows
especially in wadis subject to flash-flooding, but dies during droughts (peak flowering
March-April). It is common in parts of north-east Jordan, where Basalt Wheatear at least
formerly bred. Possible association between Z. spinosa and Basalt Wheatear, and the effect
of drought on both would be interesting to study. The bird's 'feeding territory' was rather
large, c.200 m x 500 m, but it spent most time in a better-vegetated depression of c.30 m * 80
m, which was excavated when a road was built. On 30 March the bird captured a scorpion,
which was broken into pieces and completely consumed.
Despite straightforward separation from superficially similar black wheatears discussed
above, the bird possessed some odd features requiring further investigation. On the right
side of the breast were two feathers with whitish tips c.l mm wide, and its wing formula
was outside the previously known variation of warriae (Fig. 10, Table 1), possibly suggesting
hybridisation with O. 1. lugens ? A blood sample was taken.
O. 1. warriae appears to be an early-winter immigrant, with most records of short-
staying individuals in December, suggesting the birds move further south thereafter. This
corresponds with the theory that warriae regularly disperses longer distances like O. /.
persica (Shirihai et al. 2011) and has a longer, more pointed wing than O. 1. lugens (Shirihai
et al. 2011). Most records are from deserts or partially modified habitats, but that in January
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
232
Bull. B.O.C. 2012 132(4)
TABLE 1
Basic biometrics of Basalt Wheatear Oenanthe lugens warriae, including bird trapped
in Ovda Valley, Israel, on 22 March 2012, and the two specimens in the Hemprich
& Ehrenberg collection. Museum fur Natiirkunde, Berlin (measurements, in mm,
taken by H. Shirihai, based on standard protocols of Svensson 1992).
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2010 was in northern Israel, which has a Mediterranean climate and where even O. I lugens
is a vagrant. Most records involved young males, which age/sex class might more regularly
move longer distances.
Other records of Basalt Wheatear away from its breeding grounds
As part of ongoing study into the lugens complex, in January-April 2012 I visited the
Natural History Museum, Tring, Museum fur Naturkunde, Berlin (ZMB), Naturhistorisches
Museum Wien, American Museum of Natural History, New York, Tel Aviv University
Zoological Museum, and Jordan Natural Museum, Yarmouk University. Two overlooked
specimens of warriae were identified in Berlin: one from 'Nubia' (see below) originally
labelled leucura (but more recently placed with specimens of leucopyga), and one from
Sinai, Egypt, found among the O. picata but labelled leucopyga (Figs. 15-20). The first had
already been brought to my attention by F. D. Steinheimer, while investigating potentially
older available names for warriae. I compared their biometrics with all available material
of warriae and, with the help of S. Frahnert & P. Eckhoff, retrieved the following data
concerning these specimens from the museum archive:
ZMB 4851: immature, collected by W. Hemprich & C. G. Ehrenberg [in October 1823]
in Tor [El-Tor, Egypt]. It belongs to Hemprich & Ehrenberg's eighth shipment (Lichtenstein
Legend to figures on page opposite
Figures 15-20. The two overlooked Basalt Wheatear Oenanthe lugens warriae skins from the Hemprich &
Ehrenberg collection, held in Berlin (ZMB 4854 and ZMB 4851) (H. Shirihai). Fig. 18 shows the original label
of ZMB 4854, and Figs. 19-20 those of ZMB 4851.
Figures 21-22. First-summer male Basalt Wheatear Oenanthe lugens warriae, near Urfa, south-east Turkey,
May 2011 (E. Yogurtcuoglu). Note the predominantly black body, clearly short outermost primary (pi) vs.
the tip of the primary-coverts and p2, which tends to fall level with p5.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
233
Bull. B.O.C. 2012 132(4)
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
234
Bull. B.O.C. 2012 132(4)
1824) sent from Alexandria on 16 November 1823 to Berlin (arrived 6 May 1824). Three
specimens (nos. 247-249) of Saxicola leucura capito nigro are mentioned (Lichtenstein 1824).
One (ZMB 4851) remained at ZMB, two were given away. Given the date of the shipment,
the specimens should have been collected in July-October 1823. The date was further
restricted based on the specimen's plumage.
ZMB 4854: immature, collected by W. Hemprich & C. G. Ehrenberg [in November 1821]
in 'Nubien' [route along the Nile between first and second cataract, near Aswan/Egypt].
From Hemprich & Ehrenberg' s seventh shipment (Lichtenstein 1823), sent from Alexandria
on 1 October 1822 and arrived in Berlin on 19 March 1823. Twelve specimens (no. 426-437)
of Saxicola leucura capito nigro were received taken in October / November at Deram and
Gornu [localities unknown] (Lichtenstein 1823). Three (ZMB 4852-4854) remained in Berlin
while nine were given away. ZMB 4852 and ZMB 4853 are juveniles of O. leucopyga. The
collecting date of ZMB 4854 was further restricted based on the specimen's plumage.
Biometrics appear in Table 1. Identified as warriae by the broad black tail-band and
smaller size, including shorter and narrower bill (unlike juvenile White-crowned Wheatear,
and Black Wheatear). Unlike both of the latter species and male opistholeuca, both have more
pointed wings, longer primary projection, but relatively shorter tail and broader tail-band,
as well as the diagnostic white bases to the inner webs of the regimes.
No date is available for either specimen, but plumage and wear best fit first-years,
taken in November-January for the bird from Egypt / Sudan, and October-December for
the Sinai bird. No sex is given, and individual variation and bleaching renders any attempt
to sex them unreliable. They further prove that warriae can migrate to Africa and how Basalt
Wheatear has confused ornithologists from the outset.
Basalt Wheatear in Turkey
A Basalt Wheatear found in south-east Turkey, in May 2011 (Figs. 21-22), was first
reported by D. Domuschiev (www.spatiawildlife.com/en/News/birding-Turkey-Basalt-
new-subspecies.html), but because it was paired with a Finsch's Wheatear some observers
speculated that the bird was a black morph of the latter species (www.surfbirds.com/
forum/showpost.php?p=32140&postcount=7). This hybridisation event was confirmed by
the birds' close pair-bond (E. F. Henriksen, K. M. Olsen, E. Yogurtcuoglu pers. comm.), and
both sexes were observed feeding two young that successfully fledged (D. Domuschiev
pers. comm.). The pair involved a male Basalt and female Finsch's Wheatears.
The male was definitely not a O.finschii for which there is no evidence of a black morph:
c.20 photographs of the Turkish bird reveal that it had the typical plumage and structure
of warriae, and sound-recordings clearly show that its song was unlike Finsch's. Aside from
the predominantly black plumage, the wing formula best fits Basalt: pi fell clearly short of
the tip of the primary-coverts (in Finsch's it tends to extend beyond this), and p2 is level
with p5, whereas in Finsch's it is closer to p6. The bird was a first-summer based on moult
limits and the much-reduced whitish-grey flashes in the remiges.
This record is of some interest. O. /. warriae nests only in southern Syria and north-
east Jordan, suggesting that this bird was an overshooting migrant, c.500 km north of its
usual breeding grounds. Dispersal and hybridisation perhaps reflect the apparently severe
decrease in this wheatear's population. In north-east Jordan, in spring 2012, 1 was unable to
find any, and a decline has already been postulated, due to a series of drought years in the
basalt desert (Khoury et al. 2010). Previous claims of hybridisation with O. 1. lugens could
be genuine and reflective of climatic conditions, but also correspond to the close genetic
relationship between the lugens complex (including warriae ) and finschii (Aliabadian et al.
2012; M. Schweizer & HS work in progress).
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Hadoram Shirihai
235
Bull. B.O.C. 2012 132(4)
Acknowledgements
I thank the many friends mentioned, especially Ehud Dovrat, who discussed with me the 1982 record and
translated his field notes pertaining to that bird. Uri Makover, Yoav Perlman and James P. Smith kindly sent
their field notes concerning the 2004 and 2010 records, and Amir Balaban, Amir Ben Dove, Itai Berger, Danile
Berko wic, Dick Forsman, Barak Granit, Yoav Perlman and Itai Shanni shared my field observations in March
2012. Rafael Armada, Alon Beer, David Jefferson, Uri Makover, Magnus Ullman, Yoav Perlman and Emin
Yogurtcuoglu provided their photographs. Eyal Shohat of the Israeli Rarity and Distribution Committee
confirmed their re-evaluation of the two Eilat records. Dobromir Domuschiev, Eva F. Henriksen, Klaus
Mailing and Emin Yogurtcuoglu also provided information on the Turkish record. Guy Kirwan made several
suggestions that improved the manuscript. Staff at the Natural History Museum, Tring, and American
Museum of Natural History, New York, are thanked for their assistance. Dr F. D. Steinheimer kindly checked
nomenclatural issues, and Dr Sylke Frahnert and Pascal Eckhoff at ZMB were exceedingly helpful, especially
in providing historical data. Lars Svensson has encouraged my wheatear studies throughout and made
several helpful comments that improved the final draft. Manuel Schweizer provided early genetic results
from our research into Oenanthe and useful suggestions concerning the manuscript. The referees, Pierre-
Andre Crochet, Jose Luis Copete and Magnus Ullman, made several valuable comments.
References:
Aliabadian, M., Kaboli, M., Forschler, M. I., Nijman, V., Chamani, A., Tillier, A., Prodon, R., Pasquet, E.,
Ericson, P. G. P. & Zuccon, D. 2012. Convergent evolution of morphological and ecological traits in the
open-habitat chat complex (Aves, Muscicapidae: Saxicolinae). Mol. Phyl. & Evol. 65: 35-45.
Khoury, F., Fdrschler, M. I., Janaydeh. M., Aliabadian, M. & Al-Hmoud, A.-R. 2010. Distribution, habitat
and differentiation of the poorly-known black morph of Mourning Wheatear Oenanthe lugens in Jordan.
Sandgrouse 32: 113-119.
Lichtenstein, H. 1823. Verzeichniss der siebenten Sndung der Herren Doctoren Hemprich und Ehrenber,
welche vom Januar bis Julius 1822 in Ober-Aegypten und Nubien gesammelt und am 19. Marz 1823
angekommen ist. Unpubl. ms in Museum fur Naturkunde an der Humboldt-Universitat zu Berlin,
Historische Bild- u. Schriftgutsammlungen, Bestand: Zool. Mus., Signatur ZMB S I, Hemprich &
Ehrenberg, III: 75-110.
Lichtenstein, H. 1824. Verzeichniss der Gegenstande, welche von den Doctoren Ehrenberg und Hemprich in
den und in ihrer achten Sendung aus Arabien und Agypten iibersandt worden und am 6. Mai 1824 im
Museum angekommen sind. Unpubl. ms in Museum fur Naturkunde an der Humboldt-Universitat zu
Berlin, Historische Bild- u. Schriftgutsammlungen, Bestand: Zool. Mus., Signatur ZMB S I, Hemprich
& Ehrenberg, III: 111-180.
Shirihai, H. 1996. The birds of Israel. Academic Press, London.
Shirihai, H. 1999. Fifty species new to Israel, 1979-1998: their discovery and documentation, with tips on
identification. Sandgrouse 21: 45-105.
Shirihai, H. & Svensson, L. in prep. Handbook of Western Palearctic birds. Christopher Helm, London.
Shirihai, H., Keijl, G., Gargallo, G. & Copete, J. L. in prep. Wheatears. Christopher Helm, London.
Shirihai, H., Kirwan, G. M. & Helbig, A. J. 2011. A new taxon in the Mourning Wheatear Oenanthe lugens
complex. Bull. Brit. Orn. Cl. 131: 270-291.
Svensson, L. 1992. Identification guide to European passerines. Fourth edn. Privately published, Stockholm.
Address: c/o Ausserdorfstrafie 6, 8052 Zurich, Switzerland, e-mail: albatross_shirihai@hotmail.com
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
236
Bull. B.O.C. 2012 132(4)
The collecting history and distribution of Dusky
Friarbird Philemon fuscicapillus (Morotai, Indonesia)
elucidated from museum data
by Ludovic Besson
Received 26 March 2012
Summary.™ A comprehensive search of ornithological collections throughout the
world yielded a total of 75 specimens of Dusky Friarbird Philemon fuscicapillus, a
Vulnerable species, held in 25 museums. This represents 48 additional specimens
to those already known. The history of this material is discussed. Five collectors
were identified: C. Allen, H. A. Bernstein, J. M. Dumas, A. Hueting and G. A. L.
Haan. The 18 collected by Charles Allen on behalf of Wallace are syntypes. All
70 documented specimens are from Morotai. There is no historical proof of the
presence of the species on Halmahera or Bacan, despite that these islands are
frequently considered part of the species' range.
Natural history collections are important sources of data on avian distribution and
ecology (Green & Scharlemann 2003). For some poorly known species, museum data,
often unpublished (Collar & Rudyanto 2003), can yield precise details (Haffer 2003). The
genus Philemon Vieillot, 1816, comprises 16 species (Higgins et al. 2008) found in Wallacea,
New Guinea, the Bismarcks, Australia and New Caledonia. One of the seven species in
Wallacea, Dusky Friarbird P. fuscicapillus (Wallace, 1862) is the only Philemon considered
globally threatened, listed as Vulnerable (www.birdlife.org/datazone/speciesfactsheet.
php?id=5319). An enigmatic species (BirdLife International 2001), knowledge even of its
distribution is incomplete, although since Gray (1869) P. fuscicapillus has been generally
presumed to occur on three of the northern Moluccas, namely Morotai, Halmahera and
Bacan. However, as noted by Stattersfield & Capper (2000), BirdLife International (2001,
2004) and Higgins et al. (2008) historical data are available only from the first island, recent
observations from the second (Fuller s.d., Poulsen & Lambert 2000, Jahraus 2005, Dashper
2009) and there is no real evidence that it occurs on the third (Hartert 1903, Lambert 1994).
N. J. Collar, the principal author of BirdLife International (2001), examined 26 specimens.
Stattersfield & Capper (2000), BirdLife International (2001, 2004) and Higgins et al. (2008)
specify that the bird's ecology is little known. An additional problem is that Dusky Friarbird
can be confused with the mimetic Dusky -brown Oriole Oriolus phaeochromus G. R. Gray, 1861
(Diamond 1982); photographs of both species are available at http://orientalbirdimages.org/.
My objective was to use specimens and other archives to clarify the collecting history and
distribution of this friarbird.
Material and Methods
The following abbreviations are used. AMNH: American Museum of Natural History,
New York (USA); ANSP: Academy of Natural Sciences of Philadelphia (USA); BMNH:
Natural History Museum, Tring (UK); BOUM: Museum d'Histoire Naturelle de Bourges
(France); CU: Cornell University, Ithaca (USA); CUMZ: University Museum of Zoology,
Cambridge (UK); eBEAC: electronic Bulletin for European Avian Curators; FMNH: Field
Museum of Natural History, Chicago (USA); GBIF: Global Biodiversity Information
Facility; ICOM: International Council of Museums; LivCM: Liverpool Museum (UK);
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
237
Bull. B.O.C. 2012 132(4)
MCZ: Museum of Comparative Zoology, Harvard (USA); MH: Museum Heineanum,
Halberstadt (Germany); MHNMON: Museum d'Histoire Naturelle de Montauban (France);
MZPW: Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw (Poland);
MNHN: Museum National d'Histoire Naturelle, Paris (France); MZB: Museum Zoologicum
Bogoriense, Java (Indonesia); NCB Naturalis: Netherlands Centre for Biodiversity Naturalis,
Leiden (Netherlands); NHColl-L: Natural History Collections Listserver; NMNH: National
Museum of Natural History, Washington DC (USA); NMW: Naturhistorisches Museum,
Wien (Austria); NRM: Naturhistoriska Riksmuseet, Stockholm (Sweden); OB: Oriental
Birding; ORNIS: Ornithology Information System; OUM: Oxford University Museum (UK);
RMNH*: Rijksmuseum van Natuurlijke Historie, Leiden (Netherlands); SDM: Darwin
Museum, Moscow (Russia); SMF: Senckenberg Museum Frankfurt (Germany); SNMBG:
Staatliches Naturhistorisches Museum, Braunschweig (Germany); UMB: Uberseemuseum,
Bremen (Germany); YIO: Yamashina Institute for Ornithology, Tokyo (Japan); ZMA*:
Zoologisch Museum, University of Amsterdam (Netherlands); ZMB: Museum fur
Naturkunde, Zentralinstitut der Humboldt-Universitat, Berlin (Germany). * = all now held
at Naturalis (NCB). Specimens are referred to by their current registration number. Names
of islands follow White & Bruce (1986).
Search for specimens.— This commenced at BMNH because this is where Alfred Russel
Wallace, who described the species, sent his collections (Baker 2001). Online databases
(e.g., GBIF and ORNIS) were then consulted. To widen the search, 423 institutions on five
continents were contacted, using Wood et al. (1982), White & Bruce (1986), Wood & Schnell
(1986), Mearns & Mearns (1998), Roselaar (2003) and ICOM. Finally calls for contributions
were issued on internet fora (NHCOLL-L, eBEAC, OB). All details can be obtained on
request from the author.
Documentation. — Collecting data are no longer available for many specimens and
information on their labels is often insufficient or fragmented (Fisher & Warr 2003).
Consequently, for such a little-known species as Dusky Friarbird it was necessary to make
the best possible use of all specimens. Where possible, the labels on each specimen were
compared with collection records to avoid problems of the type mentioned by Rasmussen
& Prys-Jones (2003). Conversely, the few published facts (localities, dates, collectors) were
verified using the specimens.
Curators often recognise specimens from certain collectors by their preparation
style (M. Portas & C. Fisher pers. comm.) but very little has been published on this,
usually in respect to a few specific cases, e.g., John Gilbert (Fisher 1992). This method of
identification must be used cautiously as collectors were numerous at the end of the 19th
century and could produce similar specimens despite some variation. However, in dealing
with specimens of the same species, as here, direct observation or photographs permit
similarities in preparation to be readily appreciated. Given that the specimens come from a
well-defined region worked by a small number of collectors, such comparison is pertinent.
Knowledge of the collector's identity enables us to access relevant historical records (Fisher
& Warr 2003) and to determine the collecting date.
As preparation technique provides just one indication of the collector's identity, it was
always used in conjunction with other data such as the specimen's origin, handwriting
and terms used on the labels, or the specimen's date of entry into the relevant collection.
Descriptions were made from specimens for which the collector is clearly authenticated by
the label. Birds mounted and reprepared as flat skins are not included, as this leaves no
indication of the original method of preparation.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
238
Bull. B.O.C. 2012 132(4)
Results
In addition to the information available in the GBIF and ORNIS databases, I was
able to check for the presence of Dusky Friarbird specimens in 272 collections of the 423
contacted. Seventy-five specimens were found in 25 institutions, of which photographs of
63 were obtained. The specimens occur as follows: Europe 58 (Austria: three, France: ten,
Germany: seven, Netherlands: 20, Poland, Russia and Sweden: one each, UK: 15); North
America 11 (all USA); and Asia six (Indonesia: five, Japan: one). They were collected in
1860, 1861, 1898, 1912, 1938 and 1950. Seventy were collected on Morotai and five possess no
locality or collector information. Of the 75, 56 are clearly linked to one of the following five
collectors: A. R. Wallace, H. A. Bernstein, J. M. Dumas, A. Hueting and G. A. L. de Haan. No
specimens exist from Halmahera or Bacan. The material comprises 49 skins and 26 stuffed
birds (eight of them reprepared as fiat skins). No tissues appear to have been preserved
(Wood et al. 1982, Wood & Snell 1986; pers. obs.) or eggs. The only accessible bones are
the skulls and limbs of specimens. A few notes on gonadal development are available, but
nothing concerning stomach contents.
Discussion
Full details of the Dusky Friarbird specimens are presented in Table 1. The total number
of specimens found (75) is relatively small for an extant species.
Co Hectors.— Ornithological exploration of the Moluccas is well known (Junge 1953,
White & Bruce 1986). Only three collectors appear to have worked on Morotai: Charles
Allen, Heinrich A. Bernstein and Gunter A. L. de Haan. Lendon (1946), Christian (1950)
and missions by the FAO in 1975-81, mentioned by White & Bruce (1986), did not collect
specimens. The Snellius Expedition 1929-30 (van Riel 1937) studied only oceanic, not
terrestrial, fauna. However, information provided by specimens in the present study,
permits the addition of three names: Joannes Maximiliaan Dumas, Anton Hueting and,
curiously, Alfred R. Wallace. Indeed, Wallace visited Halmahera in March and September
1858, and again in 1860 for a few days without collecting, but never Morotai (Wallace 1862,
1869). The species' first recorded collector is thus Allen, Wallace's young assistant who he
mentioned by name. Allen's collections on Morotai were made from mid-October 1860 to
mid-March 1861 (Baker 1995, 2001). Being their true owner, Wallace labelled them himself.
Wallace (1869) considered the species endemic to Morotai.
Bernstein, a doctor on Java, made nine expeditions in 1860-65, following a route
through the Malay archipelago very similar to that of Wallace, and sent 4,645 Indonesian
birds to Leiden (Finsch & Martens 1866, Wallace 1869, Jansen 2008). Details are given by
Jansen (2008). Some errors are associated with his Morotai friarbirds. Firstly, the collection
date of a female, 14 November 1861, cannot be correct as on this date he was on Halmahera.
It was perhaps taken on 14 December 1861, on which date he definitely was on Morotai
(corroborated by other specimens). Secondly, a mounted specimen is clearly dated 3
September 1862, yet Bernstein was far to the south on Obi from 17 July to 2 September 1862
and on Bacan on 9-18 September 1862 (dates corroborated by specimens taken, for example,
on Obi on 27 August 1862 and Bacan on 11 September: Hoek Ostende et al. 1997: 32, 187).
Consequently, 3 September 1861 is much more plausible, as his presence on Morotai is
corroborated by other specimens taken there on the same day. Nothing suggests that the
locality is incorrect. Thirdly, a transcription error exists for BMNH 1879.4.5.1380 (collected
29 September 1861). This date is clearly given on the specimen's label, but was transcribed
22 September by Gadow (1884: 279). None of Bernstein's Dusky Friarbird specimens is from
Halmahera or Bacan, despite frequent visits to these islands.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
239
Bull. B.O.C. 2012 132(4)
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Bull. B.O.C. 2012 132(4)
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Ludovic Besson
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Mount H. A. Bernstein ? 13 Sep 1861 Morotai
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14 Dec 1861?
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Institution plus Method of Date of Institution, collection. Field collector and Age Sex Collection Locality Status
registration no., where conservation accession naturalist merchant and/or collecting number date
available previous number
Ludovic Besson
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Ludovic Besson
243
Bull. B.O.C. 2012 132(4)
Dumas accompanied Alfred H. Everett's visit to the Moluccas (Sharpe 1906, LeCroy
& Jansen 2011). When Everett died, Dumas continued, visiting several islands including
Burn and Morotai, but not Halmahera or Bacan, and his Dusky Friarbirds were exclusively
collected on Morotai towards the end of 1898 (LeCroy & Jansen 2011). He also took at least
two entomological specimens on the island in November 1898 (Rothschild 1908: 174).
The Dutch missionary and ethnographer Hueting worked around Tobelo, Halmahera,
between 7 August 1896 and 1934. Though he lived mainly on Halmahera and the only
specimen of the friarbird that he collected (in 1912) was one of a batch of 97 skins of 37
species from 'Halmahera' sent to RMNH, it originated from Morotai (based on the label).
The Dutch collector de Haan made several expeditions in the Moluccas (van Bemmel
& Voous 1953, White & Bruce 1986), in 1938 and around 1950. His friarbird specimens are
very well documented and are all from Morotai; they were collected during both periods.
The five collectors mentioned above are probably the only ones to have visited Morotai
(Junge 1953, White & Bruce 1986, Fransen et al. 1997, Hoek Ostende et al. 1997, Dekker 2003,
Dekker & Quaisser 2006; AMNH, BMNH, FMNH, GBIF, MCZ, MNHN, ORNIS, USNM and
YIO online databases). None of the specimens was collected on Halmahera, which island all
of these collectors explored, with the notable exception of Dumas (LeCroy & Jansen 2011),
or on Bacan. Similarly, collections made on Halmahera and Bacan by other collectors (Gray
1860, Guillemard 1885, Nehrkorn 1894, Berlepsch 1901, Hartert 1903, Kuroda 1938, 1939,
Kuroda 1957, Ripley 1959) did not yield the species, and Hartert (1903) clearly stated that
no specimens were collected on Bacan.
The skins and their labels.— Observation, either directly or from photographs, of 48 of
the 49 specimens, permits their classification into five batches of homogeneous appearance,
clearly associated with each of the five collectors above. Preparation (Fig. 1) and labels (Fig.
2) have permitted clarification or confirmation of the collectors of several Dusky Friarbird
specimens. It is highly improbable that any specimen is from any island other than Morotai.
Of course, this possibility cannot be entirely excluded as collections could have disappeared
since 1860 (see below), but it is very unlikely that specimens taken elsewhere over the
course of a century by five collectors of different nationalities would all have disappeared.
Allen's skins have their feet crossed and tied, and their preparation is relatively poor.
The bird's neck is often unnaturally long, the bill set almost in line with the body and the
feathers are displaced (Fig. 1A). They were stuffed in such a way that the breast bulges
and the abdomen is very flat when viewed in profile. Wallace labelled them himself, but
was not their actual collector: the ten pre-printed labels completed by hand (Baker 2001)
are marked 'Collected by A. R. Wallace. 186.' or 'Collected for A. R. Wallace. 186.' (thereby
demonstrating that he employed others). The handwritten data indicates only the year, 1860
or 1861, an English transcription of the locality, 'Morty Is.' or 'Morty Island', and sometimes
the sex. Two are numbered, probably by Allen (Fig. 2W). Another of Allen's specimens
(SNMB 3629), clearly taken on Morotai, confirms that it was taken in 1860-61. AMNH
697224 and OUM 15207 are unlabelled and the institutions lack any other data that might
trace their provenance. Although both specimens strongly resemble Allen's preparations,
without deeper analysis (e.g.. X-ray) it can be said only that he was probably their collector.
Bernstein' s carefully prepared specimens are characterised by a wooden stick in the body's
axis (Fig. 3). Their silhouette is quite slender and they have been stuffed homogeneously
making them appear almost fresh (Fig. IB). In all of the specimens definitely collected by
Bernstein and preserved as flat skins the stick is not obvious, meaning the feathers must
be moved to see it (Fig. 3; MNHN 1873.359). Marks on the tarsi bear witness to the cut-off
part of the stick to which the feet were originally attached, probably until arrival in Europe.
Only Bernstein's skins possess this feature, which, together with their overall appearance.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
244
Bull. B.O.C. 2012 132(4)
A
B
UMZC 27/Mel/28/j/l MNHN 1873 359
USNM 178136
D
Ha
RMNH 134524 ZMA 48201
Figure 1. Comparison of Dusky Friarbird Philemon fuscicapillus specimens taken by the five collectors.
A: Charles Allen skin, face and profile (Michael Brooke, © Cambridge University Museum of Zoology);
B: Bernstein skin, face and profile (Ludovic Besson, © Museum National d'FIistoire Naturelle, Paris); D:
Dumas skin, face and profile (James Dean, © National Museum of Natural History, Washington DC); Hu:
Hueting skin, face and profile, (Eelco Kruidenier, © NCB Naturalis); Ha: de Haan skin, face (Tineke Prins, ©
Zooldgisch Museum, University of Amsterdam).
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
245
Bull. B.O.C. 2012 132(4)
Figure 2. Examples of labels attached to
Dusky Friarbird Philemon fuscicapillus
specimens. W: Wallace (Hein van Grouw,
© Natural History Museum, Tring); B:
Bernstein, back and front (Ludovic Besson,
© Museum National d'Histoire Naturelle,
Paris); D: Dumas (Hein van Grouw, ©
Natural History Museum, Tring); Ha: de
Haan (Tineke Prins, © Zoologisch Museum,
University of Amsterdam); F: Finsch (James
Dean, © National Museum of Natural
History, Washington DC).
permits their certain identification.
His handwritten labels, often with a
black border on one side, regularly
feature the name of the island in
Dutch ('Morotai'), the precise date
(day, month and year) and the sex.
Occasionally, the number of the box
in which it was sent to Europe ('Kish
in Dutch) also appears (Fig. 2B). MH
3091 lacks a label. It was mentioned
in 1882 by Heine & Reichenow but
arrived at MH before then meaning
that Dumas, Hueting or de Haan can
be excluded. Its general appearance
and the presence of a stick identify
Bernstein as the collector. In three
skins the wooden rod is present, but
there are very few data associated
with them and the collector's name
is missing. Two are strongly linked
to the Netherlands: MNHN 1875.356
was given to Paris by Schlegel, the
director of RMNH, in 1875 (Fig. 4) and
ZMA 25377 arrived in Amsterdam
before 1894; both were collected on
Morotai. MNHN 2000.1252 came via
Boucard. It lacks a date or locality,
but must have been taken before
1894 (Anon. 1905). All three were
obviously prepared by the same
person. Apart from the wooden
rod, they are identical to Bernstein's
specimens (Fig. 3), but doubt remains
concerning their collection date.
w
U< <
i yii <
BMNH 1969.30.17
B
Virrtaux
/Wv ;
MNHN 1873.359
D
t .
£jft ^ Pa c n.fA pff o_
A if <r - / /
9-d
BMNH 1900.2.8.18
$ ■ : ■
ZMA 48200
dcJ>,
<<}****>. ft*.
USNM 178136
5 cm
Bernstein visited Morotai again in 1863, which explains why their preparation is slightly
different from that of the others.
Dumas' skins are easily recognised, appearing fat and thickset, the head 'sunk' into
the shoulders so that the neck seems to disappear. Sometimes there is a thread through
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
246
Bull. B.O.C. 2012 132(4)
MNHN 1873.358
MNHN 1873.359 MNHN
Figure 3. The six Museum National d'Histoire Naturelle, Paris, specimens in ventral view with, from left
to right, two Bernstein specimens (stick broken), two Bernstein specimens (stick intact) and two Wallace
specimens (Ludovic Besson, © Museum National d'Histoire Naturelle, Paris)
the nostrils holding the mandibles together. The feet are crossed and tied (Fig. ID). They
are sometimes labelled 'Philemon fuscicapilla' , with the locality 'Morty Id' in characteristic
handwriting (Fig. 2D), although BMNH 1969.41.1217 is erroneously transcribed 'Phileomon
fuscicapilla' . BMNH 1969.41.1216 and OUM 12482 are thickset and are labelled only 'Morty
Id'. This label, the handwriting, presence of string binding the beak and general appearance
identifies these specimens as being taken by Dumas. For CU 9482, only its preparation
permits this attribution as the label is lost, but Dumas was probably the collector.
Hueting's specimen preparation resembles that of Bernstein (Fig. lHu) but he collected
just one Dusky Friarbird, the label of which is missing.
De Haan's specimens are prepared with a wooden stick through the axis of the body
and a thread in the nostrils holding the mandibles together (ZMA 48200; http://ip30.eti.uva.
nl/zma3d/detail.php?id=410&sort=alphabet&type=family). They are similar to Bernstein's
specimens except that the shoulders are more pronounced and the feet rest parallel to the
stick, rather than being crossed over it (Fig. lHa). Labelling is precise and, for the ZMA
specimens, indicate age (Fig. 2Ha), gonad state and overall length.
© 2012 The Authors; lournal compilation © 2012 British Ornithologists' Club
Ludovic Besson
247
Bull. B.O.C. 2012 132(4)
Figure 4. Extract from the Catalogue General des Entrees Mammiferes-Oiseaux, vol. 5, showing the donation
of MNHN 1875.356 by Prof. Schlegel (Ludovic Besson, © Museum National d'Histoire Naturelle, Paris)
Specimen trading.— Dispersion of collections, through sale or exchange, was frequent
in the 19th and early 20th centuries. In 1863, an advertisement in Ibis offered Dusky
Friarbirds from the Wallace collection for 12 shillings. Specimens transited many traders
and both public and private collections. Repeated trading often resulted in transcription
errors. For example, a batch of 35 birds partly collected by Allen bears the number
1862.9.11,1-35, not 1869.2.11,1-35 as indicated by Sharpe (1906) (Baker 2001). With respect
to BMNH 1900.2.8.18-19, Dumas, identified on their labels with the wrong initial 'G' (FI.
van Grouw pers. comm.), is their collector. Although Sharpe (1906: 285, 361) indicated these
specimens as having been collected by Everett, the latter died on 18 June 1898 (Anon. 1898)
and therefore could not have visited Morotai at the end of that year.
Trade in material often resulted in labels being replaced and original data being
retranscribed. Otto Finsch did this conscientiously (Rasmussen & Prys-Jones 2003) with
the labels of MCZ 39598 and USNM 178136 exact copies of RMNH specimens (Fig. 2F;
handwriting authenticated by H. van Grouw). In contrast, it is well known that the
Verreaux brothers noted collecting data with very little accuracy (Mearns & Mearns 1998,
Rasmussen & Prys-Jones 2003). In the present case, the Maison Verreaux obtained only
Bernstein specimens, which are now at MNHN. They bear the collector's original label with
the stamp of Maison Verreaux (Fig. 2B), removing any ambiguity. Another specimen from
this supplier, perhaps obtained by Wallace and formerly in the Turin museum (Salvadori
1881), is currently lost (M. Calvini & G. Doria pers. comm.). Although Boucard (1876)
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
248
Bull. B.O.C. 2012 132(4)
mentioned that he had duplicate specimens of this friarbird from 'Batchian', none of the
six specimens of his that I have located are definitely from this island; five originate from
Morotai and the provenance of the other is not specifically stated.
Unlabelled specimens.— Of the eight specimens, some are mounted but others have
been reprepared as flat skins. BOUM 12.Mel.31 (http://orientalbirdimages.org/search.
php?Bird_ID=1355) is from Albert Maes (1846-1914), a member of the French Zoological
Society from 1893 (Blanchard & Caullery 1915, Schaeck 1919), who assembled 8,000 birds
with elements from most Indonesian islands. None of his specimens bears any indication
of being collected by Hueting, who can be excluded, and de Haan was not of the same era.
Other collectors must be considered, as some of their specimens passed through institutions
and persons who exchanged with Maes.
AMNH 8079 is identified by the original scientific name, Tropidorhynchus fuscicapillus.
This nomenclature was also used by Bernstein before Gray removed the species to Philemon
in 1869, and suggests an early origin for this specimen, although the collector cannot be
identified with any certainty.
ANSP 18469 formed part of a very early collection and is labelled only with the species'
original name and the locality, 'Morotai'. A second label states ' Microphilemon fuscicapillus' ,
a genus proposed by Mathews (1912), which also appears on the labels of NMW specimens
taken by Bernstein. Use of Morotai (Wallace preferred 'Morty') and their handwriting
suggest Bernstein, but this is unconfirmed.
NRM 570190 reached Stockholm in 1866 according to its label, but the museum register
indicates 1864 (U. Johansson pers. comm.), and MNHN 1864.350 and 1866.400 arrived in
1864 and 1866, and MHNMON-O-1361 arrived in 1876. These dates mean they could not
have been collected by Dumas, Hueting or de Haan. The most likely provenance for MNHN
1866.400 is Bernstein because it was exchanged with Maison Verreaux (see above). ZMB
2000.13881 lacks any information.
Syntypes.— Wallace's (1862, 1869) collections were made prior to 1862, and included
the friarbirds procured Allen from which he described P. fuscicapillus. Following Art.
73.2 of the Code (ICZN 1999), they must be considered syntypes of Tropidorhynchus
fuscicapillus Wallace, 1862, because Wallace did not select a single type. Their status does
not appear to have been stated on the specimens, and has therefore gone unnoticed in
some cases. Interestingly, several published lists of types mention Wallace as their collector
(Gadow 1884, Sharpe 1906) whereas Warren & Harrison (1971) named Allen for BMNH
1873.5.12.1010 and the red (syntype) labels on BMNH 1862.9.11.22-23 also specify Allen.
Conclusion
This inventory identified 18 syntypes of Dusky Friarbird, with two other specimens
probably sharing this status. This list is still open as it has proved impossible to establish
the total number of specimens collected by Allen and analysed by Wallace. Several errors
have been corrected in archives and publications, collectors specified for some specimens,
and a more thorough analysis might permit reevalutation of the specimens which have no
data. Study of specimens of both sexes and all ages demonstrates that the few available
illustrations in the literature could lead to incorrect field identifications and confusion
with the species' mimic. Dusky-brown Oriole. Finally, the available data confirm that
Dusky Friarbird is endemic to Morotai, with no evidence of its presence on the islands of
Halmahera or Bacan.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Ludovic Besson
249
Bull. B.O.C. 2012 132(4)
Acknowledgements
I thank all of the institutions who replied to my requests, particularly the following and their personnel with
whom exchanges were always interesting and fruitful: AMNH: Mary LeCroy, Thomas J. Trombone; ANSP:
Nathan H. Rice; BMNH: Hein van Grouw; CU: Kimberly Bostwick, Charles M. Dardia; MCZ: Jeremiah
Trimble; LivCM: Clem Fisher; MH: Bernd Nicolai; MHNMON: Aude Bergeret, Olivier Duchein, Edmee
Ladier; MZPW: Dariusz Iwan; MNHN: Patrick Bousses, Marie J. Portas, Claire Voisin; MZB: Mohammad
Irham; NCB Naturalis (RMNH and ZMA): Justin J. F. J. Jansen, Steven van der Mije, Tineke Prins, Kees
Roselaar; NMNH: James Dean; NMW: Ernst Bauernfeind; NRM: Goran Frisk, Ulf Johansson; OUM: Malgosia
Nowak-Kemp; SDM: Igor V. Fadeev; SMF: Gerald Mayr; SNMBG: Bettina Borges-Naumer, Michaela
Forthuber; UMB: Peter-Rene Becker; CUMZ: Michael Brooke; YIO: Takeshi Yamasaki; ZMB: Sylke Frahnert.
N. J. Collar and J. Diamond shared their knowledge of Wallacea, Rosemarie Filippi (BOUM) translated
German texts, Laurent Nourrit translated Italian texts, Jean-Fran^ois Voisin (MNHN) and Frank A. Maddi
read the manuscript, and Edith Cotin translated the text into English. Finally, I thank my referees Colin
Trainor and Frank Lambert, and the editor G. M. Kirwan, for their advice and corrections.
References:
Anon. 1898. Obituary [Alfred H. Everett]. Ibis 4(4): 627-628.
Anon. 1905. Death of M. Adolphe Boucard, C.M.Z.S. Ibis 8(5): 299-300.
Baker, D. B. 1995. Pfeiffer, Wallace, Allen and Smith: the discovery of the Hymenoptera of the Malay
archipelago. Archiv. Nat. Hist. 23: 153-200.
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BESSON@ville-bourges.fr
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Full tree resolution of Polyplectron Temminck, 1813,
confirms species status of Hainan P. katsumatae
Rothschild, 1906, and Bornean Peacock-Pheasants
P. schleiermacheri Briiggemann, 1877
by G. W. H. Davison, Chang Jiang, Zhang Zhengwang & Chen De
Received 2 April 2012
Summary. — Additional molecular (cy tochrome-fr, D-loop and OvoG) sequences have
permitted resolution of a phylogenetic tree for all eight taxa of peacock-pheasants
Polyplectron Temminck, 1813. This confirms previous work indicating that Hainan
Peacock-Pheasant Polyplectron katsumatae Rothschild, 1906, and Bornean Peacock-
Pheasant P. schleiermacheri Briiggemann, 1877, are well-differentiated species,
a conclusion further supported by plumage, morphology and, in the case of P.
schleiermacheri, vocalisations. The tree confirms that P. katsumatae is one of a cluster
of species geographically peripheral to, and derived from, a widespread mainland
Asian ancestor that is today represented by P. bicalcaratum, and that these two
species show greater genetic divergence than exists between Grey P. bicalcaratum
and Sumatran P. chalcurum or Mountain Peacock-Pheasants P. inopinatum.
For more than a decade there has been renewed interest in the taxonomy (Zheng 2002,
Madge & McGowan 2002, Penhallurick & Walters 2005), nomenclature (Dickinson 2001)
and systematics (Kimball et al. 2001) of taxa in the genus of peacock-pheasants Polyplectron
Temminck, 1813. At species level two key questions have been whether the taxon katsumatae
Rothschild, 1906, from Hainan is specifically distinct from Chinese mainland Grey Peacock-
Pheasant P. bicalcaratum (Linnaeus, 1758) and whether schleiermacheri Briiggemann, 1877,
from Borneo is specifically distinct from Malaysian Peacock-Pheasant P. malacense (Scopoli,
1786) of the Malay Peninsula.
Bornean Peacock-Pheasant P. schleiermacheri was accepted as a species by Beebe
(1922), Peters (1934), Inskipp et al (1996), Johnsgard (1986, 1999), Sibley & Monroe (1990),
MacKinnon & Phillipps (1993), McGowan (1994), Smythies (1999), Dickinson (2003), Myers
(2009) and Phillipps & Phillips (2009). It was reduced to a subspecies of P. malacense by
Delacour (1951), and this was accepted by Smythies (1957, 1981) and Hennache & Ottaviani
(2006).
Hainan Peacock-Pheasant P. katsumatae was first described as a species, and authors
who have retained it at this rank include Beebe (1922), Inskipp et al. (1996), MacKinnon &
Phillipps (2000), Madge & McGowan (2002), Zheng (2002) and Liang & Zhang (2011). Peters
(1934) was the first author to treat it as a subspecies of P. bicalcaratum, while those who have
maintained it at subspecies level include Delacour (1951), Meyer de Schauensee (1984),
Johnsgard (1986, 1999), Cheng (1987), Sibley & Monroe (1990), Gao (1991, 1999), McGowan
(1994), Li (1996, 2004), Dickinson (2003) and Hennache & Ottaviani (2006). Chang et al.
(2008), who obtained cytochrome-^ and OvoG sequences from nine individual katsumatae
and six bicalcaratum, found the differences sufficient to recognise them as species and, more
significantly, found that they are not each other's closest relatives, being interpolated in
phylogenetic estimates by Mountain Peacock-Pheasant P. inopinatum (Rothschild, 1903)
from the Malay Peninsula.
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Collar (2009) critiqued the paper by Chang et al. (2008), re-examined the type series
of P. katsumatae in the American Museum of Natural History, New York (AMNH) and
supported species-level recognition for this taxon based on criteria advocated by Tobias et
al. (2010). He found it difficult to accept that P. bicalcaratum could be more closely related to
the biogeographically distant P. inopinatum than to the 'adjacent' P. katsumatae. Collar (2009)
also found it difficult to accept that the strongly distinct P. inopinatum is phylogenetically
interpolated between P. bicalcaratum and P. katsumatae because the two latter taxa are more
similar in their plumage. Kimball et al. (2001), who sampled mitochondrial DNA from six
Polyplectron taxa but not P. katsumatae or P. schleiermacheri, found that Sumatran Peacock-
Pheasant P. chalcurum is sister to P. bicalcaratum, and P. inopinatum is sister to these two.
Like Collar (2009) they considered the phylogeny incongruent with the geographical
implications, but lack of sampling of P. chalcurum by Chang et al. (2008) and of P. katsumatae
by Kimball et al. (2001) make direct comparison of their results impossible. Another potential
criticism of Chang et al. (2008) could be their failure to sample extensively across the range
of mainland Asian P. bicalcaratum, as would be desirable to ensure that P. katsumatae is not
nested within divergent mainland haplotypes of that species.
Our aims in this study were to resolve the species status and the closest relatives of
katsumatae and schleiermacheri, and to determine the validity of doubts raised by Collar
(2009) on the compatibility of molecular and morphological information.
Methods
We obtained fresh material of P. katsumatae and P. schleiermacheri (see
Acknowledgements). The material of P. katsumatae was from the feather illustrated by Lee
et al. (2005: PI. 5). The material of P. schleiermacheri was from eggshell membranes, feathers
and wet tissue of captives in Singapore, which showed no history or signs of hybridisation
(L. K. C. Kuah pers. comm.). From AMNH we obtained wet tissue from a sample of P.
bicalcaratum independent of those analysed by Chang et al. (2008), as well as a wet tissue
sample of Crimson-headed Partridge Haematortyx sanguiniceps. We used the primers listed
by Kimball et al. (1999) and Armstrong et al. (2001). We applied a DTT-based adaptation
of the QIAGEN filter kit for extracting DNA from toepads and feathers of other taxa
sampled via the Natural History Museum, Tring (BMNH), AMNH and Raffles Museum
of Biodiversity Research, National University of Singapore (RMBR), including Painted
Spurfowl Galloperdix lunulata, Germain's Peacock-Pheasant P. germaini, P. chalcurum and
P. inopinatum. Cytochrom e-b, D-loop and OvoG sequences were obtained by amplification,
sequencing and overlapping partial sequences using the software SeqEdit (Applied
Biosystems, USA). OvoG sequences (not obtained for P. katsumatae and Haematortyx) were
aligned with Clustal X, this and other procedures being concordant with those used by
Chang et al. (2008). We used these sequences to re-analyse a complete phylogenetic tree for
all eight Polyplectron taxa, adding the GenBank information from the papers by Kimball et al.
(2001) and Chang et al. (2008) for Polyplectron species, as well as cytochrom e-b, D-loop and
OvoG sequences from a range of taxa as outgroups. Phylogenetic trees were constructed
using the maximum likelihood (ML) method implemented in PAUP* 4.0bl0 (Swofford 1998)
and a Bayesian tree was constructed using MrBayes software (http://mrbayes.sourceforge.
net) at allcompat and halfcompat settings.
Like Collar (2009), we examined specimens of P. katsumatae in AMNH, and took notes
on plumage and measurements. Because ocellus colours vary with angle of incident light,
we viewed them all with the skin placed between the observer and light source, facing left.
We supplemented these with notes on posture, plumage and bare-parts coloration from
photographs of several live birds of both sexes in Li (2004) and Corder (2001), taken both
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
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Bull. B.O.C. 2012 132(4)
in the wild and at the South China Institute for Endangered Animals, Guangdong, China,
by Y.-R. Gao.
We took notes on behaviour, plumage and voice of live captive P. schleiermacheri,
and compared sonograms with closely related species, but were unable to obtain sound-
recordings of P. katsumatae. Sonograms were prepared using Avisoft Sonagraph Pro
developed by Raimund Specht, Berlin, with a software sampling rate of 16,000 Hz 16 bits,
using a Soundblaster Audio Card (Creative Labs Inc) on an IBM compatible PC running
Microsoft Windows.
Results
DNA.—Our results, shown in Figs. 1-2, were in full accord with those of Kimball et al.
(2001) and Chang et al. (2008). Like Kimball et al. (2001) we found that P. chalcurum and P.
bicalcaratum are sister taxa, and P. inopinatum is sister to this pair. Like Chang et al. (2008) we
found that P. katsumatae is the next closest and that P. germaini is basal to all these preceding
taxa. Our new material of P. bicalcaratum, P. chalcurum, P. inopinatum, P. katsumatae and P.
germaini clustered respectively with the previously sequenced materials from each of these
taxa. Although we had only one additional specimen of P. bicalcaratum, independent of
those used by Kimball et al. (2001) and Chang et al. (2008), there is no evidence that any of
these taxa (in particular mainland P. bicalcaratum vs. Hainan P. katsumatae) are paraphyletic.
We found that P. schleiermacheri was sister to P. malacense, and concurred with Kimball
et al. (2001) and Chang et al. (2008) that Palawan Peacock-Pheasant P. napoleonis is basal to
all congeners.
P. inopinatum
P. bicalcaratum
P. bicalcaratum wet tissue
P. chalcurum
P. katsumatae DJ
P. katsumatae Z
P. germaini
P. malacense
P. schleiermacheri egg D J
P, schleiermacheri feather D
P. schleiermacheri toepad
P. napoleonis
Figure 1. Consensus tree showing maximum likelihood (ML) values using all non -Polyplectron taxa as the
outgroup and employing 50% majority rule bootstrap.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
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Bull. B.O.C. 2012 132(4)
— Haematortyx sanguiniceps
P. inopinatum
~ P. bicalcaratum
L P. bicalcaratum wet tissue
P. chalcurum
P. katsumatae
L P. katsumatae feather
~ P. germaini
— P. malacense
~ P. schleiermacheri egg membrane
P schleiermacheri feather
L P schleiermacheri toepad
. P napoleonis
Alectoris chukar
I l 0.050 expected changes per site
Figure 2. Phylogenetic tree created using MrBayes at halfcompat setting, rooted with Gallus, with branch
length proportional to expected base changes per site.
Depending on the tree root, Haematortyx may be most closely related to Polyplectron
(Fig. 2). We obtained short sequences from Galloperdix lunulata, too fragmentary for use
in phylogenetic reconstruction and not differing from sequences in Polyplectron. All three
genera have multiple spurs, and male G. lunulata has faint iridescence on the wing-coverts
and tail, as well as white ocelli. Employment of different outgroups did not affect the
topology of the Polyplectron clade, with the highly ornamented P. napoleonis always basal to
other Polyplectron.
Plumage. — Vive distinctions between P. katsumatae and P. bicalcaratum were listed by
Collar (2009): remarkably smaller size; steel green (not steel purplish) ocelli on the wings
and mantle; shorter crown feathers; crown darker than (not paler or uniform with) the
neck and mantle; and darker body plumage as a result of much denser vermiculations.
In addition we noted that the bare facial skin of live P. katsumatae is brilliant red, and this
colour extends well behind the eye, and onto the mandible and maxilla as far as the anterior
tip of the nares. The red colour extends over the operculum and can meet across the ridge
of the culmen. The divide between bare facial skin and feathered supercilium is distinct
and complete, whereas in P. bicalcaratum the facial skin bears numerous tiny feathers and
there is very little bare skin posterior to the eye. In male P. bicalcaratum the bare skin ranges
from dirty grey to pale yellow or buff, and extends to the base of the mandible but not the
maxilla. Even if the colour variations in the bare skin of P. bicalcaratum are associated with
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
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Bull. B.O.C. 2012 132(4)
reproduction or hormonal change, the colour
never reaches the intense red shown by P.
katsumatae. Thus the colour, shape, extent
and degree of bareness of the facial skin are
all distinct. We found that one live female P.
katsumatae had bare red facial skin and pale
irides, whereas in female P. bicalcaratum the
facial skin is sparsely feathered, grey to pale
yellow or pinkish, and the irides brown.
We concur with Collar (2009) that the
ocelli on the wings and mantle are steel green
not steel purplish; the distinction is not subtle
and it sorts all males of P. katsumatae from all
those of P. bicalcaratum. In addition we note
that the ocelli on the wings of P. katsumatae
are broader than long and have a tiny distal
point or Tail' extending over the shaft. In
both taxa the ocelli on the uppertail-coverts
and rectrices are green, but they are brighter
and greener in P. katsumatae. The ocelli on the
rectrices of P. katsumatae are smaller, 17-21
mm max. length measured parallel to the
shaft on the central rectrix ( n = 4), compared
with 24-27 mm in P. bicalcaratum ( n = 4) and 19-21 mm in P. germaini ( n = 4). The overall
effect is of smaller ocelli, further from the tip of the tail (Table 1) but the blurred outline of
the ocelli makes measurements imprecise.
In male P. bicalcaratum the ocelli on the inner and outer webs are separate on at least the
inner 4-5 median tail-coverts. In male P. katsumatae the ocelli are fused at the shaft on all the
median tail-coverts. Our impression is that more pairs of rectrices of male P. bicalcaratum
have the ocelli separate (on 5-6 central pairs, and sometimes on all 10-12 pairs of rectrices),
and this is true of fewer pairs of rectrices in male P. katsumatae (ocelli separate on 3-4 central
pairs, never on all 9-10 pairs of rectrices) but that this is subject to much variation.
Collar (2009) noted that in body plumage P. katsumatae 'is somewhat darker, as a result
of much denser vermiculations than bicalcaratum' . We add that these vermiculations are
smaller, neater, rounder and more orderly, and less vermiform. We also found that both
sexes are substantially browner, less grey, especially on the wings. Collar (2009) noted that
the crown of P. katsumatae is darker, but we add that there is no iridescence on the crown
or nape, a further distinction from P. bicalcaratum , which has purple iridescence on those
feathers. In some features P. katsumatae resembles P. germaini (small size, red facial skin,
darker and browner plumage) but in others it does not (pale iris, pale surrounds to ocelli).
In P. schleiermacheri, numerous major plumage distinctions from P. malacense have
already been described and illustrated by Beebe (1922), Delacour (1951), Hennache &
Ottaviani (2006) and others, including the short crest, pure white chin, white central breast
stripe, black breast and flanks, iridescence at the breast-sides, transverse subterminal bars
on the rectrices, and brilliant orange-red facial skin.
Voice. — Sonograms showed that the call of P. schleiermacheri differs from those of P.
napoleonis and P. malacense. The call of P. schleiermacheri is an explosive high-pitched clarion
with an almost quacking quality, usually consisting of two notes, kank-kank, but sometimes
one kank per call, comprising broad frequency spectrum noise between 1.5 and 5.0 kHz,
TABLE 1
Ocellus dimensions in three taxa of Polyplectron.
Each measurement refers to the maximum length
of one ocellus on the outer web of a central rectrix
of an individual male.
Length of ocellus
(mm)
Distance to tip of
rectrix (mm)
Ratio
P. katsumatae
17
56
3.294
18
61
3.389
21
64
3.048
21
broken
-
P. bicalcaratum
24
61
2.542
24
67
2.792
26
65
2.500
27
56
2.074
P. germaini
19
58
3.052
21
65
3.095
21
48
2.286
21
62
2.952
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G. W. H. Davison et al.
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Bull. B.O.C. 2012 132(4)
kHz
2 *
0.2 0.4 0.6 sec
i *
1.0 1.2 sec
Figure 3 (upper left). Sonogram and pressure recording of two-note kank-kank advertising call of male
Polyplectron schleiermacheri in captivity (GWHD: 20050618); (upper right) pressure recording of seven in
a long series of single-note advertising calls of male P. napoleonis in captivity (GWHD: 20071104); (lower
left) sonogram of single-note tchorr advertising call of male P. malacense, Pasoh, Negeri Sembilan, Malaysia
(K. W. Scriven: 1969); (lower right) sonogram of long clucking series by male P. malacense that eventually
degenerates to single tchorr calls, the series continuing for several minutes, Pasoh, Negeri Sembilan, Malaysia
(K. W. Scriven: 1969).
with max. energy output around 3.5 kHz. Each note lasts c.0.15 seconds, and the duration
of the two-note call is c.0.4 seconds. These calls are given every few seconds in a long series.
Male P. napoleonis give higher pitched, more staccato, broad frequency spectrum single
notes, each lasting c.0.15 seconds and given every few seconds in a long series at intervals
of c.2 seconds. Max. energy output of 15-20 /iPa is at a frequency around 3.5 kHz.
In addition to its soft, mellow, deliberate two-note pure whistle, male P. malacense gives
single harsh notes (the tchorr call: Medway & Wells 1976); each note is a broad spectrum
noise with max. energy output at around 2.5 kHz, with duration of 0.5 seconds (Fig. 3).
Such notes are given at long intervals, but males sometimes deliver a long series of notes
beginning with a single explosive tchorr tailing away into a prolonged chain of decelerating
clucks (Fig. 3, lower right), repeated with progressively fewer clucks and greater emphasis
on the first note at each repetition, until reaching a long series of spaced, single tchorr notes
(Fig. 3, lower left). Such a progression in calling is unknown in P. schleiermacheri or P.
napoleonis, but this could be due to shortage of field observations.
We could not be sure of comparing like with like, as the advertising kank-kank of P.
schleiermacheri is distinct from both the advertising mellow whistle and harsh alarm tchorr
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of P. malacense, but in captive P. schleiermacheri the call was used in aggressive and sexual
contexts not alarm. There could be additional calls not yet known, but we could be sure
that the known calls of P. schleiermacheri, P. napoleonis and P. malacense are all very different.
Discussion
There is now ample evidence from DNA sequencing and morphology that P. katsumatae,
and from DNA sequencing, morphology and voice that P. schleiermacheri, are recognisable at
species level. We have shown that there are numerous plumage and bare-parts distinctions
between katsumatae and bicalcaratum additional to those listed by Collar (2009). The strong
differences in facial skin colour and shape, ocellus colour and proportions that we noted in
specimens are also clearly evident in photographs in Corder (2001), Li (2004) and Liang &
Zhang (2011). The cumulative evidence from Kimball et al. (2001), Chang et al. (2008) and
the present study shows that both taxa fully meet the Phylogenetic as well as the Biological
Species Concept (BSC), using criteria described by Helbig et al. (2002), Parkin et al. (2006)
and Tobias et al. (2010). Using the scoring system of Tobias et al. (2010), Collar (2009) scored
P. katsumatae with seven out of a potential nine points, but the taxon might have scored
even higher if a different set of morphological characters (e.g., the bare-part coloration less
obvious in specimens) had been taken into account.
We cannot demonstrate failure to hybridise as a criterion for applying the BSC, as
each taxon is allopatric in the wild and many Galliformes hybridise freely in captivity
with intergeneric and even some interfamilial hybrids known (McCarthy 2006). But
correlations between bare-part coloration and display features of Polyplectron are important
in each species (Davison 1983); and the displays of P. chalcurum and P. inopinatum, which
are located between P. bicalcaratum and P. katsumatae in the tree, are distinct in posture,
movement and signal type (Davison 1985, 1992). We interpret this as strong evidence that
each taxon studied here meets BSC criteria.
The finding by Chang et al. (2008) that P. katsumatae is not the sister taxon of P.
bicalcaratum is additionally supported here, as both P. chalcurum and P. inopinatum show
greater DNA sequence similarity to P. bicalcaratum than any of the three does to P. katsumatae.
This is powerful evidence that P. katsumatae merits species rank. In contrast to Kimball et al.
(2001) and Collar (2009) we do not consider this to be inconsistent with geography. On the
contrary, expansion of an ancestral Asian mainland 'bicalcaratum' down the western margin
of the Sunda Shelf, during a drier period around the Pliocene-Pleistocene transition, would
have been followed by the isolation of several peripheral populations caused by sea-level
rise and higher rainfall. Speciation events were timed by Kimball et al. (2001) at 0.7 (0.2-1. 2)
MYA for P. chalcurum, 1.5 (0.8-2.2) MYA for P. inopinatum, and by Chang et al. (2008) at
1.4 (1.1-1. 7) MYA for P. katsumatae. The broad margins of error, and possible differences
in the rates of accumulation of differences in DNA sequences, are sufficient to allow for
apparent discrepancies. For example, one would expect Sumatran P. chalcurum to have been
isolated prior to or at the same time as Malay Peninsula P. inopinatum, and the margins of
error given by Kimball et al. (2001) permit this. The picture is of a once more widespread
mainland Asian taxon, from which peripheral populations were calved by one or a series
of near-contemporaneous events, possibly speciation of P. germaini having been slightly the
earliest and resulting in the form that was geographically least likely to mix with others
(chalcurum, inopinatum, katsumatae) after isolation.
The first separation of Hainan from the mainland of southern China was c.2 MYA
(Zhang 1999), followed by several episodes of unification during low sea-level glacials and
separation during high sea-level interglacials. Pollen data from Hainan and mainland China
(Zheng 2000, Lin & Zhang 2001) suggest the types of vegetation change these populations
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G. W. H. Davison et al.
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would have experienced. Glacial events were accompanied by changes in rainfall on
Hainan with the expansion of grassland, restriction of Dacrydium, Castanopsis and Quercus
to upland forest, and increased abundance of Mallotus and Casuarina in the lowlands during
drier glacial maxima (Zheng 2000), a pattern opposite to that on the mainland for which at
least the Last Glacial Maximum was wetter.
This Polyplectron pattern is consistent with isolation of peripheral populations of
other mainland Asian birds in western landmasses of the Sunda Shelf, e.g. Crested Argus
Rheinardict ocellata, Hill Prinia Prinia atrogularis and Black-throated Sunbird Aethopyga saturata
in the Malay Peninsula, and Grey-headed Woodpecker Picus canus in the Malay Peninsula
and Sumatra. It is not our purpose to put a case for the dating of range extensions by these
birds, merely to point out biogeographical parallels. We envisage that for Polyplectron such
events would have occurred when other lowland Polyplectron were confined to a moister
eastern or central block of the Sunda Shelf, P. napoleonis, P. schleiermacheri and P. malacense
representing successive differentiation of populations from east to west and isolating P.
inopinatum in the mountains of the Malay Peninsula as wetter conditions permitted the
spread of P. malacense and the retreat of ' bicalcar atum' in the lowlands.
Acknowledgements
We thank Lee Kwok Shin and Bosco Chan of the Kadoorie Farm and Botanic Garden, Hong Kong, for a
fresh feather sample of P. katsumatae, J. Corder for material of P. inopinatum, and Paul Sweet (American
Museum of Natural History, New York), Mark Adams (Tring, BMNH) and Wang Luan Keng (Raffles
Museum of Biodiversity Research, RMBR) for permission to extract DNA material from other species. We
thank Lawrence K. C. Kuah for access to fresh genetic material of P. schleiermacheri and for permission to
make observations and sound-recordings. Gerald Neo of the Agri-Food and Veterinary Authority (AVA),
Singapore, facilitated CITES and import permits for various samples. Additional material was extracted
and sequenced by Jagdish Kaur Chahlil (University Malaysia Sabah), with advice and tree construction
by Menno Schilthuizen and Dick Groenenberg (Naturalis, Leiden). Sound-recordings of P. schleiermacheri
and P. napoleonis were analysed by Mike McGuire, Queensland, while those of P. malacense derive from
sonograms provided to GWHD by Joan Hall-Craggs in 1976, based on recordings by K. W. Scriven. We
greatly appreciate the support and encouragement of R. W. Stein, and D. R. Wells, University of Cambridge
Zoology Museum, and we acknowledge the role of the late Gao Yuren, formerly of the South China Institute
for Endangered Animals, in having maintained a stock of P. katsumatae for study today.
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Taxonomy and nomenclature of the Stonechat complex
Saxicola torquatus sensu lato in the Caspian region
by Lars Svensson, Hadoram Shirihai, Sylke Frahnert & Edward C. Dickinson
Received 20 April 2012
Summary.— We discuss the taxonomy of the Stonechat, the accepted broad species
Saxicola torquatus , and find convincing reasons for recognising three species:
European Stonechat S. rubicola , Eastern Stonechat S. maurus and African Stonechat
S. torquatus. The nomenclature of the taxa breeding in the Caspian region is
revisited and, based on an analysis of the original type descriptions and all relevant
literature, and of four preserved specimens of Ehrenberg's taxon hemprichii from
1833 now in Berlin, we conclude that the name variegatus should not be applied to
the taxon breeding north of the Caucasus but instead to the population in eastern
Turkey and Transcaucasia, present-day armenicus. This places armenicus in the
synonymy of variegatus, the latter having priority. Thirdly, a name is required for
the north Caspian population. The name hemprichii is the oldest available and valid
name for this population.
Background
Several important contributions have recently been published regarding the taxonomy
and nomenclature of the broad Stonechat complex Saxicola torquatus, of which a few have
foscused on those taxa in the Caspian region. It is now generally agreed that the complex is
best treated as (at least) three different species, as first suggested by Wittmann et al. (1995)
and Wink et al. in Urquhart (2002). More recent studies have supported this based on more
extensive genetic sampling (Illera et al. 2008, Zink et al. 2009, 2010). Other than molecular
indicators, the split is backed by consistent morphological differences and the fact that two
of the three groups breed in partial sympatry apparently without interbreeding. In the area
of sympatry, clear differences in habitat selection are evident. Genetically, Fuerteventura
Stonechat S. dacotiae is more closely related to the European group of taxa of S. torquatus
sensu lato than this group is related to similar-looking Asian taxa, a fact that has lent
further support to the split. Retaining a large single Stonechat species, and avoiding a non-
monophyletic species, would require that the quite distinct Fuerteventura Stonechat be
subsumed as a subspecies under the Stonechat, hardly a desirable path.
The division into three species gives the following taxonomy:
1. The mainly European form becomes the European Stonechat Saxicola rubicola (von
Linne, 1766). Two subspecies, connected by intermediate populations, are warranted:
rubicola in much of continental Europe reaching east to the Caucasus and Transcaucasia,
and hibernans (E. Hartert, 1910), breeding in the British Isles and coastal western France,
north-west Spain and Portugal; breeders in the western Netherlands might also best be
referred to this race.
2. The mainly Asian form becomes the Eastern Stonechat S. maurus (Pallas, 1773). This
species has several recognised subspecies, but of immediate concern here are the
following three Palearctic taxa: maurus breeding in Russia and marginally in adjacent
parts of eastern Europe, in western and south-central Siberia, Kazakhstan and parts
of Transcaspia, east to north-west Mongolia and south to Afghanistan and northern
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Pakistan (possibly also in north-east Iran); variegatus (S. G. Gmelin, 1774) in eastern
Ukraine on the lower Don, east Crimea, the Kalmykiya plains north-east to the Volga
Delta area, in the south on the northern slopes of Caucasus, apparently reaching north
Azerbaijan (hereafter 'NCT' for 'North Caspian Taxon'); and armenicus Stegmann, 1935,
in north-east Turkey, Caucasus (apparently at least locally on the southern slopes),
Transcaucasia and western and southern Iran (hereafter 'SCT' for 'South Caspian
Taxon'). Status in Azerbaijan is not well known and needs to be better established. See
Fig. 1 for an overview of the distribution of the discussed taxa.
3. The mainly African form, breeding in much of Africa with c.15 subspecies described
(some of which might warrant separation as species when better studied), but also with
a population in south-west Saudi Arabia and western Yemen, becomes the African
Stonechat S. torquatus (von Linne, 1766). The only race of any concern in a Palearctic
context (and, depending on Palearctic limits adopted, arguably not even then) is
southern Arabian felix G. L. Bates, 1936.
In preparing a new handbook to the birds of the Western Palearctic (Shirihai & Svensson
in prep.) all available specimens of relevant taxa in several major collections (Tring, New
York, Leiden, Paris, Berlin, Bonn, Stockholm, Copenhagen and Moscow) were examined
by LS, and two field trips were undertaken by LS to north-east Turkey, the Caucasus and
Transcaucasia in order, among other aims, to clarify the taxonomy of these stonechats. HS
& LS also met in Tring and examined the collection there together. Finally, for the purpose
of this work, HS & SF examined and measured two of the Ehrenberg types of S. maurus
hemprichii in the Berlin museum. SF further searched and discovered two more types of this
taxon in Berlin, as discussed below.
Figure 1. Map of the breeding range in the Caspian region of the discussed subspecies within the Stonechat
complex, here treated as European Stonechat Saxicola rubicola (blue outline) and Eastern Stonechat S. maurus
(yellow). The two taxa of the latter breeding in the Caspian region, current day variegatus and armenicus, have
here been labelled as 'NCT' (= north Caspian taxon; orange) and 'SCT' (= south Caspian taxon; light blue),
respectively, the reason for which is explained in the text (after Vaurie 1959, Cramp 1988, Urquhart 2002,
Collar 2005, Zink et al. 2009; M. Banik in litt. 2011).
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Eastern Stonechats of the Caspian region
As mentioned above there are distinct populations of S, maurus in the north and south
Caucasus. The correct application of names to these populations will be discussed after
further background is given, but these two subspecies can be circumscribed as follows:
NCT.— The northern population has a very characteristic male plumage with extensive
white portions on each side of the inner tail (between half and three-quarters of the outer
tail feathers white), not unlike the pattern in many wheatears or male Red-backed Shrike
Lanius collurio. This is easily seen on a flying bird, but can be more difficult to confirm on
perched birds with closed tails. The amount of white in the tail on males is subject to a
subtle cline; at its maximum in the Volga Delta region, becoming slightly more restricted
in the south. Both sexes resemble nominate maurus from further east and north in that they
have a large unstreaked pale rump patch, huffish when fresh, white when abraded and
bleached. In comparison, European Stonechat S. rubicola has a streaked rump with usually
limited white. The NCT male has a concentrated red-brown chest patch and a large white
patch on the neck-sides, again more like nominate maurus and different from most rubicola.
The NCT female is very similar to females of both nominate maurus and the SCT, and
usually can be identified only if handled. The female has much less white on the base of the
rec trices than the male, generally requiring that the tail-coverts be lifted to reveal the white,
and a few females lack any. The name used for the NCT in Vaurie (1959), Ripley (1964),
Urquhart (2002) and Dickinson (2003) has been variegatus.
SCT.— Resembles the NCT but differs in somewhat larger size (a certain overlap in
all measurements between the two; Table 1), in having less white on the tail base in male
plumage (at most the inner half of feathers white, but more commonly only one-third),
often not visible even in flight, and darker, more saturated colours. In particular the dark
TABLE 1
Biometrics (mm) of four taxa of the Stonechat complex Saxicola torquatus sensu lato , nomenclature
according to conclusions presented here. Measurements taken from skins according to standards
described in Svensson (1992). Data separated by sex and presented as range, sample size (n), mean (in
bold) and standard deviation (SD). All measurements by LS.
Taxon
Saxicola r. rubicola
8. maurus variegatus
'South Caspian Taxon'
S. maurus hemprichii
'North Caspian Taxon'
Saxicola m. maurus
Wing (max.)
8 63.0-69.0 mm
n = 26, 66.2, SD 1.59
$ 63.0-66.5 mm
n = 17, 64.8, SD 0.93
8 70.5-80.0 mm
n = 94, 75.0, SD 1.90
$ 70.0-76.5 mm
n = 43, 72.8, SD 1.62
8 68.0-76.0 mm
n = 64, 72.0, SD 1.52
$ 66.0-72.5 mm
n = 22, 69.4, SD 1.86
8 63.0-72.0 mm
n = 22, 68.3, SD 1.97
$ 64.0-70.0 mm
n = 13, 66.7, SD 1.39
Tail
8 43.0-49.0 mm
n = 26, 46.0, SD 1.83
$ 44.0-48.5 mm
n = 17, 45.7, SD 1.26
8 47.0-56.0 mm
n 93, 51.7, SD 1.92
$ 47.0-54.5 mm
n 43, 50.2, SD 1.65
8 44.0-52.0 mm
n = 64, 49.1, SD 1.57
? 45.0-51 mm
n = 22, 48.2, SD 1.58
8 44.0-51.0 mm
n = 21, 47.5, SD 2.18
§ 45.0-50.0 mm
n = 12, 47.0, SD 1.49
Tail/wing
ratio
8 n = 26, 69.6, SD 2.41
$ n = 17, 70.5, SD 1.57
8 n = 94, 69.0, SD 1.90
? n = 43, 68.9, SD 2.16
8 n = 64, 68.2, SD 1.98
$ n = 22, 69.4, SD 1.52
8 n = 22, 69.5, SD 1.97
$ n = 12, 70.4, SD 2.21
Tarsus
8 21.0-22.9 mm
n = 25, 22.0, SD 0.60
$ 20.7-22.7 mm
n = 17, 21.9, SD 0.45
8 20.0-23.0 mm
n = 79, 21.4, SD 0.68
$ 19.5-22.2 mm
n = 41, 21.1, SD 0.65
8 19.4-22.2 mm
n = 58, 21.1, SD 0.66
$ 19.5-21.8 mm
n = 22, 20.8, SD 0.67
8 20.0-22.9 mm
n = 22, 21.2, SD 0.81
$ 19.8-22.2 mm
n = 12, 21.1, SD 0.74
Bill (to skull)
8 13.5-15.5 mm
n = 23, 14.5, SD 0.51
$ 13.1-15.3 mm
n = 16, 14.3, SD 0.64
8 13.1-15.8 mm
n = 26, 14.5, SD 0.58
$ 13.1-16.0 mm
n = 41, 14.4, SD 0.66
8 13.2-15.5 mm
n 63, 13.9, SD 0.58
$ 13.2-14.8 mm
n = 22, 13.8, SD 0.41
8 12.7-14.4 mm
n - 21, 13.9, SD 0.47
$ 13.5-14.7 mm
n = 12, 14.0, SD 0.38
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brick-red chest patch in contrast to the white belly is a striking feature on male SCT. The
chest colour corresponds to, or is a mixture of, Vandyke Red and Burnt Sienna in Ridgway
(1912, PL XIII k and II k, respectively). Females differ from the NCT in that they apparently
invariably lack white in the tail. The name used for the SCT in most modern handbooks and
checklists has been armenicus.
Samuel Gottlieb Gmelin's variegatus
In Reise durch Rufiland zur Untersuchung der drey Natur-Reiche ('Travel through Russia to
explore the three realms of nature'), pt. 3 (1774: 105-107), S. G. Gmelin described a new bird
which he found commonly in Schamachie (= present-day Samaxi in north-west Azerbaijan)
on the southern slopes of the south-east Caucasus, and further east and south en route to
Persia. He named the bird Parus variegatus (misspelled 'varietagus' in the text but obviously
a typographic error and corrected on PI. 20: 3 of the same work), gave it a fairly good
description for the time, including a detailed table of measurements, and mentioned that it
had a white tail base without being more precise. The new bird is depicted in a rather crude
woodcut illustration, but despite the picture's shortcomings one can see that it depicts a
male stonechat in autumn plumage (Gmelin travelled in August). It is believed that no
syntype survives. Gmelin collected many birds on his travels, some of which went to St.
Petersburg (but apparently none collected so early are preserved). The specimen depicted
may be considered to have been the type or one of several syntypes and may be taken to
represent the named taxon.
The identification of Gmelin's variegatus as being the NCT, i.e. with much white in the
tail in male plumage, has remained unchallenged until recently, when Mlikovsky (2011)
proposed that it referred to the SCT. His arguments were that (a) the type locality must
be regarded as Bandar-e-Anzali in Iran to where Gmelin was heading, not Samaxi where
he first saw the bird and made his detailed description; (b) since only the SCT is known to
breed in Iran this form must have been Gmelin's bird; and (c) since he read into Gmelin's
extremely brief description of the tail (literally 'in the beginning white, then black') proof
that it could only refer to the SCT. Incidentally, Mlikovsky got the tail pattern wrong when
stating that it is the distal part which is white rather than the basal. He also claimed that
Stegmann 'erroneously' believed that Gmelin's bird was described from Samaxi when the
original description indisputably states that it was first seen there, and this place has since
been correctly regarded as the type locality by Hartert (1910), Stegmann (1935), Gladkov
(1954) and Ripley (1964). Thus there is no basis for 'correcting' the type locality to Bandar-
e-Anzali, nor is there a need to do so as Samaxi (40°37'N, 48°38'E) lies within the breeding
range of the SCT (Stegmann 1935; see below). Aware that his proposals would leave the
NCT birds in need of a new name, Mlikovsky made a proposal in that context which we
will address below.
However, Mlikovsky' s conclusions matched those we had already reached for three
different reasons.
First, Gmelin provided some very detailed measurements in a table. Several of these
give rather curious distances of no practical use for today's ornithologists, but at least two
prove crucial for the identification of the new bird. These are total length and tail length.
The measurements in the table are in a variation of inches, and we owe much thanks
to P. H. Barthel (in litt. 2012) for resolving this matter. The units in the Gmelin table are
'ZolT (corresponding to the inch) and 'Linie' (corresponding to Vn of a Zoll). This means
in turn that the total length of Gmelin's bird is 140 mm and tail length 54 mm. One of us
(LS) has collected many measurements of both the NCT and the SCT, showing that the
SCT is a larger bird than the NCT, and the measurements of Gmelin's bird match only the
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TABLE 2
Known ranges for the two concerned taxa in the Caspian region compared to Gmelin's (1774)
measurements of the new bird he found. Sexes combined. Measurements from skins according to
standards in Svensson (1992), giving range, mean (in bold), sample size ( n ) and standard deviation (SD).
Total length
Tail
'North Caspian Taxon'
117-137 mm
44-52 mm
(125.8, n = 65, SD 4.75)
(48.7, n = 89, SD 1.76)
'South Caspian Taxon'
119-146 mm
47-56 mm
(131.0, n = 83, SD 5.49)
(51.2, n = 137, SD 1.97)
Gmelin's bird
140 mm
54 mm
SCT (Table 2). We are fully aware of the limitations in using Gmelin's measurements of a
freshly killed bird without knowing which detailed measuring technique he applied, and
to compare them with modern measurements of specimens. Still, the measurements are
there in the original description, they seem quite carefully taken and exact, and it would
be similarly questionable not to make use of them at all. At least they serve as supporting
though not conclusive evidence.
The second reason to believe Gmelin's bird is the SCT is rather surprisingly found in
Stegmann (1935), where it is stated that the only certain breeder of the SCT found by that
author was one collected on 20 May 1896 in Samaxi ('Schemacha'), in other words the
place where Gmelin first found his bird! How Stegmann wrote this without noticing the
apparent contradiction is hard to understand. After all, he accepted Gmelin's bird as being
the NCT with type locality in Samaxi, then goes on to name the SCT and can only report one
breeding locality for it, Samaxi. We have reasons to believe that the specimen mentioned by
Stegmann is in the Zoological Institute in St. Petersburg. Confirmation of its presence there
and its identity would naturally be of interest, but a request regarding this has met with no
success.
Finally, a third supporting
but not conclusive reason
for believing that Gmelin
described the SCT is offered by
a closer study of PI. 20: 3 of his
work, reproduced here as Fig.
2, depicting what we consider
must serve to represent the
type. Although it is impossible
to see any undisputed amount
of white on the tail base of this
bird, that could be due to the
fact that even rather large white
portions of outer rectrices
in birds are often concealed
when the tail is folded. One
might argue that there is a tiny
amount of white at the base
of the outermost rectrix, but
this could also be interpreted
as a white covert. However,
the depiction certainly shows
Figure 2. Gmelin's bird as depicted on PL 20: 3 in Gmelin (1774), here
shown to represent the type of the South Caspian Taxon of Eastern
Stonechat, now Saxicola maurus variegatus. Scanned from the original
in the library of Museum fur Naturkunde, Berlin.
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a male stonechat, possibly an adult male judging from the large white wing patch, in fresh
autumn plumage, and one that at least did not have very extensive white visible in the tail.
One might speculate that had the bird had a large amount of white (like any normal NCT
male) it would have been tempting for the author and the artist to show at least some of
this striking feature. But either the bird had very little white on the tail base, so there was
no reason to show it in the plate, or the author and artist decided it was more important
to convey the bird in an absolutely true and lifelike image concealing the white as it might
appear in life. We cannot know the answer, but we can make the educated guess that it
probably had rather limited white since none is shown.
The name to be applied to the northern population
Even before it became clear that the name variegatus had to be reassigned to the SCT,
in separating northern and southern populations we faced a problematic morphological
analysis because there appeared to be very few specimens of southern birds. For a long
time we were aware of a mere three specimens in Tring, and rather few in other museums.
However, in June 2011, HS found three full trays in Tring marked 'Saxicola torquata
hemprichii', which had previously been overlooked. These specimens, over 90 in all, proved
to be of the SCT, and immediately made it possible to compare a sufficient sample of the
southern birds with sympatric ruhicola and to be sure that the differences were not just
due to individual variation. The new material clearly showed that the two sympatric
forms differed considerably in size with no overlap in wing length (Table 1). Apart from
the size difference, they differed clearly in several morphological traits with no overlap.
Knowledgeable local ornithologists in Transcaucasia, notably V. Y. Ananian, who had
accompanied LS in Georgia, had always claimed that rubicola and the SCT behaved as
different species, but firm proof seemed to be lacking. Now both genetic and morphological
evidence supported the split.
That these Tring specimens were labelled hemprichii was a surprise because this name
lay quite deep in synonymy. The name was not mentioned in Vaurie (1959), Ripley (1964)
or Urquhart (2002). It was found in Hartert (1910: 707) as a synonym of Pratincola torquata
maura with the name variegatus senior to it in the synonymy. Use of the name hemprichii for
these specimens seems to be due to the revision by Grant & Mackworth-Praed (1947), who
provided data from Stegmann (1935) showing that males of 'armenica' were longer winged
than those associated with 'variegata' (then viewed as the northern birds) or 'maura'. Grant
& Mackworth-Praed noted that Stegmann had not discussed the name hemprichii and
wrote that Ehrenberg's description agreed, as regards the white at the base of the tail, with
armenica. Thus the Tong series' of long-winged 'armenica' came to be labelled hemprichii
based on an evaluation of Ehrenberg's brief description. The latter name, of course, has over
100 years priority over Stegmann' s name.
In fact, however, hemprichii is not what Grant & Mackworth-Praed thought. Ehrenberg
(1833) described a new stonechat that differed from rubicola by having a white base to the
tail and a white rump patch. He mentioned 20 specimens, without designating a holotype,
deposited in the Museum fur Naturkunde (ZMB), Berlin. The available type material (Figs.
3-4) shows that it has more evident white at the base of the tail and that the name cannot
apply to the SCT and must apply to the NCT, so the name on the trays of these Tring
specimens was wrong.
Mlikovsky suggested that a suitable new name for the NCT would be amaliae Buturlin,
1929. This is almost 100 years younger than the name hemprichii but more importantly it is
inapplicable as it is a synonym of rubicola, a form without any white in the tail and with a
streaked rump.
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Figure 3 (above). Three of the four Ehrenberg types
of Saxicola maurus hemprichii held in ZMB, Berlin.
The lectotype is seen at the far left, while two of the
three known paralectotypes are seen to the right of
it (H. J. Gotz © Museum fur Naturkunde, Berlin)
Figure 4 (left). One of the four Ehrenberg types of
Saxicola maurus hemprichii held in ZMB, Berlin. This
paralectotype, ZMB 4920, is preserved as a mount
(H. J. Gotz © Museum fur Naturkunde, Berlin)
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Fixation of the name Saxicola hemprichii Ehrenberg, 1833
Ehrenberg (1833) based his original description in the Symbolae physicae, found on folio
page 'aa', on 20 syntypes. He gave no locality for the species but associated it with Saxicola
Rubicola Nubiae Licht. Unfortunately Lichtenstein's name cannot be traced in printed or
unpublished catalogues (Lichtenstein 1823, 1854), or in the ornithological inventory in Berlin
or on the specimen labels. None of the latter are originals, because these were removed and
replaced by museum labels when the specimens were integrated in the collection; similar
habits existed in other European museums up to this time and perhaps later. Ehrenberg
(1833) mentioned all of the regions where they had collected as localities where they found
S. hemprichii except 'Syria' (now Lebanon; Mlikovsky & Frahnert 2011). Of the 20 specimens,
nine were registered in the museum's catalogue (started only in the 1850s and based on
label information from the available specimens) as Saxicola hemprichii, under the localities
Nubia (= Egypt / Sudan), Gumfudde (= Al Qunfudhah, Saudi Arabia), Abessinia (= Eritrea,
according to Stresemann 1954) and Egypt. Today, only four type specimens can be found in
the Berlin collection. Most of the syntypes are no longer present due to unregistered early
exchanges. Some may even have been destroyed during World War II.
These four specimens are undoubtedly the NCT being three adult males and one first-
year female, the three males with extensive white in the tail, the female with some white
on the bases of the tail feathers concealed below the tail-coverts. It is conceivable that the
other 16 specimens may have differed, especially as some females will have been less easily
identified than the specimens discussed above, and could prove, if found, to represent other
forms belonging to this species complex. It is therefore necessary to designate a lectotype.
We designate adult male ZMB 4918 (Nubia) as the lectotype, chosen because its label
shows that it was previously the only specimen labelled as 'the type' by Erwin Stresemann
(and is the only specimen so labelled). However, we have found no evidence that he
published a lectotype designation. Should such a lectotypification be found, our considered
action here will be consistent with his. Due to the lectotypification, the type locality must
be treated as Egypt / Sudan and this bird is, of course, a migrant away from its breeding
grounds in the northern Caucasus or north-west Caspian.
The three other former syntypes at ZMB, two adult males with very similar appearance
and one female, and the missing 16 further specimens, become paralectotypes.
List of types of hemprichii registered at the Museum fur Naturkunde, Berlin (extant
specimens indicated in bold; data in square brackets added from information in printed and
archive sources, or following examination of the specimens)1
Lectotype: ZMB 4918, skin, [adult] male, collected [between November 1821 and August
1822] in 'Nubien' [= Egypt / Sudan].
Paralectotype (lost): ZMB 4919, male, collected [between November 1821 and August 1822]
in 'Nubien' [= Egypt / Sudan].
Paralectotype: ZMB 4920, mount, [adult] male, collected [between November 1821 and
August 1822] in 'Nubien' [= Egypt / Sudan].
Paralectotype (lost): ZMB 4921, female, collected [between November 1821 and August
1822] in 'Nubien' [= Egypt / Sudan].
1 A further specimen of Hemprich & Ehrenberg (ZMB 4929, mount, first-year female, collected [between fuly
1823 and January 1824] in Tor [= El-Tor, Egypt] and catalogued as ' Saxicola ( Pratincola ) rubecola Bechst.', but
labelled on the mount as 'Saxicola hemprichii Ehr. 1828', was not included in the type series. As the label seems
to be younger than the catalogue entry, it is probable that Hemprich & Ehrenberg did not determine it as S.
hemprichii since it is a first-year female with scarcely visible white portions in the tail. It was determined as
S. maurus hemprichii by LS in 2012.
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Paralectotype: ZMB 4922, skin, [adult] male, collected [in February 1825] at Gumfudde
[= Al Qunfudhah, Saudi Arabia].
Paralectotype (lost): ZMB 4923, juvenile, collected [between April and July 1825] in
'Abessinien' [= Eritrea].
Paralectotype (lost): ZMB 4924, juvenile, collected [between April and July 1825] in
'Abessinien' [= Eritrea].
Paralectotype: ZMB 4925, skin, [first-year] female, collected [between April and July 1825]
in 'Abessinien' [= Eritrea].
Paralectotype (lost): ZMB 4926, juvenile, collected [between 1820 and 1825] in Egypt.
Conclusions
The above findings lead to the nomenclature, synonymy and range statements below.
SAXICOLA MAURUS
Saxicola maurus hemprichii Ehrenberg
Saxicola Hemprichii Ehrenberg, 1833, Symbolae physicae, Folio, page 'aa'. — No locality given
but associated with Nubia (in winter). Locality fixed as 'Egypt / Sudan'.
Varus Variegatus S. G. Gmelin, 1774, Reise Russl, 3, p. 105, PL 20: 3. — Shemakha (= Samaxi,
Azerbaijan). Name previously incorrectly applied to this taxon.
Range includes northern Azerbaijan, north-west Caspian shores to Volga Delta region, in
west, to east Ukraine.
Saxicola maurus variegatus S. G. Gmelin
Parus Variegatus S. G. Gmelin, 1774, Reise Russl., 3, p. 105, PI. 20: 3.— Shemakha (= Samaxi,
Azerbaijan).
Saxicola torquata armenica Stegmann, 1935, Doklady Akad. Nauk. S.S.R., n. ser., 3, 9. 47.—
Adshafana, Kurdistan.
Saxicola torquata excubitor Koelz, 1954, Contrib. Inst. Regional Explor., no 1, p. 13.— Dorud,
Luristan, Iran.
Range includes eastern Turkey, southern slopes of Caucasus, Transcaucasia, northern and
western Iran.
Acknowledgements
Thanks are due to Jose Luis Copete and David Bigas, who accompanied LS to Georgia and Armenia, and
helped greatly in the field. Thanks also to our local guides there, Giorgi Darchiashvili and Vasil Y. Ananian,
for assistance, and to the latter for sharing his insight into stonechat taxonomy. LS also thanks Andrew P.
Lassey, Andrew Grieve and Guy Kirwan who joined him in eastern Turkey in search of stonechats. We owe
thanks to Mikhail Banik for interesting new data on distribution. Sincere thanks also to the staff in various
museums who have helped access specimens and literature. In particular we would like to mention Robert
Prys-Jones, Alison Harding and Mark Adams at the Natural History Museum, Tring, Paul Sweet at the
American Museum of Natural History, New York, Ulf Johansson at Naturhistoriska Riksmuseet, Stockholm,
Claire Voisin at Museum National d'Histoire Naturelle, Paris, David Khaydarov at the Zoological Museum,
Moscow, Pascal Eckhoff at the Museum fiir Naturkunde, Berlin, and Jan Bolding Kristensen, Zoologisk
Museum, Copenhagen. Peter H. Barthel kindly helped with understanding Gmelin' s measurements.
References:
Cramp, S. (ed.) 1988. The birds of the Western Palearctic, vol. 5. Oxford Univ. Press.
Collar, N. J. 2005. Family Turdidae (thrushes). Pp. 514-807 in del Hoyo, J., Elliott, A. & Christie, D. A. (eds.)
Handbook of the birds of the world, vol. 10. Lynx Edicions, Barcelona.
Dickinson, E. C. (ed.) 2003. The Howard and Moore complete checklist of the birds of the world. Third edn.
Christopher Helm, London.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
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Ehrenberg, C. G. 1833. Symbolae physiciae seu icones et descriptions corporum naturalium novorum [...]. Pars
zoologica. Decas Prima [partim]: signatures 'y' to ' z 7 and 'aa' to 'gg'. Officina Academica, Berlin.
Gladkov, N. A. 1954. [Family Turdidae - partim]. Pp. 405-621 in Dementiev, G. P. & Gladkov, N. A. (eds.)
[Birds of the Soviet Union], vol. 6. Moscow. [In Russian.]
Gmelin, S. G. 1774. Reise durch Rufiland zur Untersuchung der drey Natur-Reiche., pt. 3, pp. 105-107 and (in sep.
vol.) PL 20: 3. Imperial Academy of Sciences, St. Petersburg.
Grant, C. H. B. & Mackworth-Praed, C. W. 1947. On the migratory Stonechats in eastern Africa. Bull. Brit.
Orn. Cl. 67: 47-48.
Hartert, E. 1910. Die Vogel der paldarktischen Fauna, Bd. 1. R. Friedlander & Sohn, Berlin.
Illera, J. C., Richardson, D. S., Helm, B., Atienza, J. C. & Emerson, B. C. 2008. Phylogenetic relationships,
biogeography and speciation in the avian genus Saxicola. Mol. Phyl. & Evol. 48: 1145-1154.
Lichtenstein, H. 1823. Verzeichnifi der Doubletten des zoologischen Museums der Konigl. Universitat zu
Berlin nebst Beschreibung vieler bisher unbekannter Arten von Saugethieren, Vogeln, Amphibien und
Fischen. I-X, 1-118. Berlin.
Lichtenstein, H. 1854. Nomenclator avium Museu zoologici berolinensis - Namenverzeichniss der in der
zoologischen Sammlung der Koniglichen Universitat zu Berlin aufgestellten Arten von Vogeln nach
den in der neueren Systematik am meisten zur Geltung gekommenen Namen und Gattungen und ihrer
Unterabtheilungen. Konigliche Akademie der Wissenschaften, Berlin: 1-123.
Mlikovsky, J. 2011. Nomenclatural and taxonomic status of birds (Aves) collected during the Gmelin
Expedition to the Caspian Sea in 1768-1774. J. Nat. Mus. ( Prague ) 180: 81-121.
Mlikovsky, J. & Frahnert, S. 2011. Type specimens and type localities of birds collected during Wilhelm
Hemprich's and Christian Ehrenberg' s expedition to Lebanon in 1824. Zootaxa 2990: 1-29.
Ripley, S. D. 1964. Turdinae. Pp. 12-177 in Peters, J. L. (ed.) Check-list of the birds of the world, vol. 10. Mus.
Comp. Zook, Cambridge, MA.
Stegmann, B. K. 1935. Zur Verbreitung und geographischen Variabilitat des Schwarzkehligen
Wiesenschmatzers. Doklady Acad. Nauk. S.S.R. n. ser. 3: 45-48.
Stresemann, E. 1954. Hemprich und Ehrenberg. Reisen zweier naturforschender Freunde im Orient
geschildert in ihren Briefen aus den Jahren 1819-1826. Abhandl. Deutschen Akad. Wissenschaften 1954(1):
1-177.
Urquhart, E. 2002. Stonechats. Christopher Helm, London.
Vaurie, C. 1959. The birds of the Palearctic fauna. Passerines. H. F. & G. Witherby, London.
Wink, M., Sauer-Gurth, H., Heidrich, P., Witt, H.-H. & Gwinner, E. 2002. A molecular phylogeny of
stonechats and related turdids. Pp. 22-30 in Urquhart, E. Stonechats. Christopher Helm, London.
Wittmann, U., Heidrich, P., Wink, M, & Gwinner, E. 1995. Speciation in the stonechat ( Saxicola torquata )
inferred from nucleotide sequences of the cytochrome-b gene. /. Zoo/. Syst. Evol. Res. 33: 116-122.
Zink, R. M., Pavlova, A., Drovetski, S., Wink, M. & Rohwer, S. 2009. Taxonomic status and evolutionary
history of the Saxicola torquata complex. Mol. Phyl. & Evol. 52: 769-773.
Zink, R. M., Pavlova, A., Drovetski, S., Wink, M. & Rohwer, S. 2010. Corrigendum to 'Taxonomic status and
evolutionary history of the Saxicola torquata complex'. Mol. Phyl. & Evol. 53: 1056-1057.
Addresses: Lars Svensson, S:ta Toras vag 28, 269 77 Torekov, Sweden, e-mail: lars@lullula.se. Hadoram
Shirihai, c/o Ausserdorfstrafie 6, 8052 Zurich, Switzerland, e-mail: albatross_shirihai@hotmail.com.
Sylke Frahnert, Museum fiir Naturkunde, Leibniz-Institut fiir Evolutions- und Biodiversitatsforschung,
Invalidenstrafie 43, D-10115 Berlin, Germany. Edward C. Dickinson, Flat 3, Bolsover Court, 19 Bolsover
Road, Eastbourne, BN20 7JG, UK.
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Further on the type series and nomenclature of the
Isabelline Shrike Lanius isabellinus
by David Pearson, Lars Svensson & Sylke Frahnert
Received 15 May 2012
Summary.— The lectotype of Lanius isabellinus at the Museum fur Naturkunde,
Berlin, has been considered to represent the race breeding in Mongolia rather than
that of the Tarim Basin region of western China, meaning that the name speculigerus
Taczanowski would become a junior synonym. However, questions have been
raised concerning this specimen, in particular regarding the small size of the
white primary patch and the 'atypical' narrow frontal extension to the black face
mask. Here we describe three further specimens from the type series (two males,
one female) recently relocated in the Berlin collection. The three type series males
(including the lectotype) vary in face markings and wing patch, but all have the
dark primaries associated with speculigerus and all can be closely matched by birds
from the Mongolian breeding grounds. The mask details and wing patch size of
the lectotype fall within the normal range of variability of the Mongolian race. One
of the other males is a very close match for a syntype of speculigerus held in Berlin.
In wing length and wing / tail ratio the type specimens accord with the Mongolian
rather than the Tarim Basin race. Plumage and structure therefore confirm that
isabellinus is the correct name for the former, and that arenarius Blyth should be
used for the latter.
Isabelline Shrike L. isabellinus was described by Hemprich & Ehrenberg (1833) from
specimens collected near Gumfude (= Al Qunfudhah), western Arabia, between 6 February
and 4 March 1825. Several taxa have since been treated as races of this species: speculigerus
Taczanowski, 1874, phoenicuroides Schalow, 1875, tsaidamensis Stegmann, 1930, and arenarius
Blyth, 1846.
Stegmann (1930) accepted four races within L. isabellinus , namely isabellinus ,
phoenicuroides, speculigerus and tsaidamensis, a treatment followed by others such as
Olivier (1945), Lefranc & Worfolk (1997), Panov (1996) and Cramp & Perrins (1997); also
by Vaurie (1959), although he placed the 'isabelline' races as a subgroup within a broad
species L. collurio (Red-backed Shrike). The races phoenicuroides and speculigerus, breeding
in Kazakhstan and Mongolia respectively, undertake long migrations to Africa and Arabia.
Both are characterised by dark flight feathers, a bold black face mask and prominent white
wing speculum in the male, and by pronounced sexual dimorphism. But whereas male
phoenicuroides has a chestnut crown contrasting with a darker brown back, a conspicuous
white supercilium and largely white underparts, male speculigerus has more uniform
isabelline-grey or sandy isabelline crown and upperparts, a buffish-tinged supercilium
and buff-washed underparts. (We are aware of certain variation, with some males having
a somewhat rufous-tinged crown, although the general upperparts coloration is still rather
uniform and paler than in phoenicuroides.)
The race regarded as nominate isabellinus by the above authors breeds in the Tarim Basin
region of Xinjiang (western China) and has a shorter migration, wintering from southern
Iran to north-west India. It resembles speculigerus in general coloration, with uniform
sandy isabelline upperparts, a poorly marked supercilium and sandy-buff underparts, but
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has pale flight feathers, typically a small or invisible white wing patch, and a duller face
mask, poorly developed in front of the eye, and it exhibits less sexual dimorphism. Race
tsaidamensis, which breeds at higher altitudes in the Tsaidam depression, Qinghai (west-
central China) is similar but larger, and shows a large white wing patch (see Panov 2009).
Russian authors such as Korelov (1970), Stepanyan (1990), Kryukov (1995), Panov
(1996, 2009) and Koblik et al. (2006) treated phoenicuroides as a full species, leaving just the
three isabelline races under L. isabellinus. It is these isabelline races that we are concerned
with here.
During their study of moult in L. isabellinus Stresemann & Stresemann (1972) recognised
five races. They reported that most birds wintering in Africa and Arabia moulted post-
migration, while those wintering in India moulted beforehand, in July-September. In the
first group they included phoenicuroides, speculigerus and nominate isabellinus ; in the second,
birds for which they used the name arenarius, and tsaidamensis. They examined a male at
the Museum fur Naturkunde in Berlin (ZMB) from Hemprich & Ehrenberg's type series
and designated this the L. isabellinus lectotype. They considered that representatives of this
race wintered in Africa, but could not identify the precise winter quarters of speculigerus.
Nor were they certain as to the breeding origin of nominate isabellinus vis-a-vis speculigerus.
The name change
One of us (Pearson 1979) compared African wintering specimens at the Natural History
Museum, Tring (BMNH), labelled L. i. isabellinus, with a series of male speculigerus from the
breeding area, kindly loaned from the Zoological Institute, St. Petersburg, by V. Loskot,
and concluded that these represented the same race. Thus, contra Stresemann & Stresemann
(1972) there appeared to be just one dark-primaried form (other than phoenicuroides)
breeding in Central Asia and migrating to Africa. Arabian wintering specimens at Tring
resembled those from Africa while, in contrast, the different moult strategy and duller,
paler appearance of Indian wintering specimens was striking. Reference to the type
description, and examination of an excellent photograph of the ZMB lectotype provided by
SF, led to the conclusion that the name isabellinus must indeed apply to the dark-primaried
African wintering race, meaning that speculigerus became a junior synonym (Pearson 2000).
Stresemann' s name arenarius was resurrected for the Xinjiang breeding race.
Panov (2009) questioned this view. He expressed doubts that the isabellinus lectotype
represents a true Mongolian speculigerus. He considered that the extension of the black loral
band narrowly across the base of the bill is atypical, noting that this pattern is frequent
in birds near the zone of hybridisation between L. isabellinus and L. collurio on the Chuya
Steppe near the Mongolian / Russian border but absent in the speculigerus terra typica of
Transbaikalia, and implied that its presence in the lectotype could indicate introgression by
collurio genes. He also considered the white speculum of this specimen to be too small for
speculigerus. We respond here to these points, give measurements of L. isabellinus museum
specimens from various breeding and non-breeding areas, and describe three isabellinus
paralectotypes at ZMB.
Analysis of the type series
In the species description, Hemprich & Ehrenberg gave as locality 'prope Gumfudam
Arabiae in Wechabitarum montibus' (near Gumfude [modern-day Al Qunfudhah, Saudi
Arabia] in the Asir Mountains in Arabia). They listed Lanius isabellinus for 'Arabia
meridionals, which means that their specimens were from south Arabia only, and would
have been included in the tenth shipment from Alexandria in November 1825 that reached
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Berlin in April 1826. In the shipment list there is an entry by Lichtenstein for 'Lanius
ruficaudus' (his name as well as Hemprich' s at that time for L. isabellinus) nos. 387-395 (six
males, four females), locality: 'Abessinien' (the locality is erroneous) (ZMB archives, Zool.
Mus., Sign. SI, Hemprich & Ehrenberg III: 191). The ZMB collection catalogue, as well as
Lichtenstein (1854), mentions four specimens of Lanius ruficaudus from Gumfude / Arabia,
which implies that the other six specimens were given away shortly after 1826. The four
specimens now in Berlin (ZMB 1887-90) as well as those given away all belong to the
type series. Hemprich and Ehrenberg arrived in Gumfude on 6 February 1825 and left on
4 March 1825 (Stresemann 1954). During that time both of them made trips into the Asir
Mountains as Stresemann described. Since no exact collecting date is recorded for each
specimen, the correct date to give is this timespan of nearly one month.
When Stresemann & Stresemann (1972) defined a lectotype for Lanius isabellinus they
had only two specimens available, a male and a female. They gave a type label to the male,
ZMB 1887. The female must have been ZMB 1888, which Meise (c.1950), in a card catalogue
of the types of the ornithological collection in Berlin, had given as available, but 'not
typical'. In 2010 we located the two missing type specimens in the Berlin collection, both
males (ZMB 1889-90). We do not know why these had been unavailable to Meise and the
Stresemanns in the mid-20th century, but this was perhaps a result of the disruption of the
collection by World War II damage. We are sure, however, that the two newly discovered
individuals certainly belong to the type series as they possess the original museum labels.
We have been unable to locate any of the remaining five or six type specimens, which
must have been given away to other museums. We briefly describe below the four type
specimens now present in the Berlin collection.
ZMB 1887. L. isabellinus lectotype. Male, 6 February-4 March 1825, Kunfuda (= Al
Qunfudhah, south-west Saudi Arabia). Upperparts pale buffy brown, tinged greyish from
nape to crown, more cinnamon on uppertail-coverts. Underparts washed pinkish-buff.
Blackish mask through ear-coverts, around eye, and over lores to bill, extending across base
of bill as frontal band c.l mm wide. Paler, ill-defined buffy supercilium. Remiges and larger
wing-coverts blackish brown, tertials and inner greater coverts with broad sandy brown
edges. Primaries 4-9 (numbered ascendently) with white bases, forming patch extending 4
mm beyond primary-coverts. Tail cinnamon-brown above, becoming slightly darker near
tip, uniformly pale cinnamon below. Wing 92 mm; tail 76 mm; second primary (p2) 1 mm
longer than p6. Fully moulted.
ZMB 1889. L. isabellinus paralectotype. Male, 6 February-4 March 1825, Kunfuda (= Al
Qunfudhah, south-west Saudi Arabia). Colour of upperparts and underparts as lectotype.
Blackish mask, reaching to bill but narrower on lores, with some dark grey (not black)
extending to forehead. Wings blacker than in lectotype, with large white primary patch
extending 12 mm beyond primary-coverts. Uppertail paler cinnamon-brown. Wing 98 mm;
tail 83.5 mm; p2 shorter than p6. Fully moulted.
Legend to figures on page opposite
Figure 1. Comparison, left to right, of ZMB 1887 ( isabellinus lectotype), ZMB 1889 ( isabellinus paralectotype),
ZMB 1990 ( isabellinus paralectotype), ZMB 58.58 and ZMB 21886 ( speculigerus syntype) (Lars Svensson, ©
Museum fur Naturkunde, Berlin)
Figure 2. Head patterns of (a) ZMB 1887 (top) and ZMB 1889, and (b) AMNH 261.414 (Lars Svensson, ©
Museum fur Naturkunde, Berlin, and American Museum of Natural History, New York)
Figure 3. Male Lanius i. isabellinus trapped at Narovlin, Kentiy, Mongolia, 27 June 2005 (Paul J. Leader)
Figure 4. Male Lanius i. isabellinus, 50 km south-west of Ulan-Ude, Transbaikalia, 30 May 2010 (Magnus
Hellstrom)
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ZMB 1890. L. isabellinus paralectotype. Male, 6 February-4 March 1825, Kunfuda (= Al
Qunfudhah, south-west Saudi Arabia). Similar to ZMB 1889, but upperparts and crown
more uniform buffy brown, and blackish mask confined to ear-coverts and spot before eye.
White primary patch extends 3 mm beyond coverts. Apparently fully moulted, but inner
six primaries browner than rest and presumably replaced earlier. Wing 99 mm; tail 86 mm;
p2 longer than p6.
ZMB 1888. L. isabellinus paralectotype. Female, 6 Feb-4 Mar 1825, Kunfuda (= Al
Qunfudhah, south-west Saudi Arabia). Buffy brown above, rather darker and browner
than lectotype, crown uniform with back. Underparts washed pale buffy brown, unbarred.
Poorly contrasting dark brown face mask. Wing feathers medium brown with pale sandy
buff tertial edges but no white primary patch. Tail and uppertail-coverts as lectotype. Wing
95 mm; tail 81 mm; p2 longer than p6.
The isabellinus lectotype has the head and body coloration, including bold black face
mask and blackish-brown primaries, typical of specimens from Mongolia and of freshly
moulted spring birds from Africa. It shows no characters suggestive of L. collurio, and
there is no indication that it is a hybrid. It is matched closely by ZMB 58.58, collected at the
Dalai Nur, Inner Mongolia, China, on 23 August 1956, which has a similarly small wing
patch and broad black loral band just reaching the edge of the culmen over the forehead.
The isabellinus paralectotype ZMB 1889, with a narrower loral band and large wing patch,
matches other examples from Africa and Mongolia, including the syntype of speculigerus
(ZMB 21886), from the Argun River, Dauria (probably now Chita Oblast, Russia), on 26
May 1873. Fig. 1 compares the three male isabellinus type specimens with ZMB 58.58 and
ZMB 21886.
Face mask and primary patch variations
The black face mask of male speculigerus is quite variable, as shown by Panov (2009:
PL 6). A frontal band such as that in the isabellinus lectotype was noted in c.25% of the
examples he detailed in his Table 1, admittedly more prevalent in certain breeding areas,
but occurring throughout most of the range. Among museum examples, we have noted
this feature in birds from Mali in January (MNHN 1966.634), Mongolia (no locality)
in June (AMNH 261.414), Kergelen River, Mongolia, in July (ZMMU 55498), and from
Dalai Nur, Mongolia, in August as mentioned above (for other museum acronyms see
Acknowledgements). Migrants from Kazakhstan in May (BMNF1 1898.9.20.624) and Kashi,
Tarim Basin, in late April (BMNH 1931.7.8.362), have also shown this trait. By contrast,
other Mongolian examples exhibit an incomplete mask, with little black on the lores, similar
to that of isabellinus paralectotype ZMB 1890. Fig. 2 shows the head pattern of two of the
ZMB type series males, one with a frontal band and one without, and of AMNFi 261.414.
Two further live examples with a black frontal band, recently photographed in Mongolia,
are illustrated in Figs. 3-4.
A white wing patch appears to be present in all males from Mongolia and all spring
males from Africa, but varies in size. That of the isabellinus lectotype is rather small, but falls
within the range of birds we have examined from Mongolia (3-10 mm beyond the primary-
coverts; n = 12) and Africa (2-10 mm; n = 30). Of the two male paralectotypes, one has a
relatively small patch, the other a large one.
Structure
The Mongolian and Tarim Basin breeding races differ slightly in structure. Table 1
compares wing and tail measurements, and second primary length of the four birds (three
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TABLE 1
Biometric comparison between the type series, breeders (A) or winterers (B) of the Mongolian race, and
spring / summer birds (C) or winterers (D) of the Tarim Basin race.
Wing (mm)
Tail (mm)
Mean T / W
no.
with p2
>p6
=p6
<p6
Type series* ( n = 4)
96.0 (92-99)
81.6 (76-86)
0.850
3
1
A (n = 13)
95.2 (92-101)
81.6 (76-87)
0.857
9
3
1
B (n = 38)
95.4 (90-102)
80.5 (75-88)
0.844
26
8
2
C(n = 17)
91.5 (89-94)
79.7 (77-83)
0.871
3
8
5
D (n = 67)
90.9 (87-95)
79.3 (75-84)
0.872
12
24
24
*As wing and tail measurements differ between the sexes by only c.l mm (cf. Cramp & Perrins 1993) we have included
the single female in the small type series.
males, one female) of the ZMB type series (taken by LS) with those of the following groups
of males from various collections: A — spring / summer birds with dark primaries from
east-central Asia (LS); B - winter / early spring birds with dark primaries from Africa and
Arabia at BMNH (DP); C — spring / summer birds with pale primaries from Afghanistan
and western Xinjiang (China) (LS); and D— winter birds with pale primaries from Pakistan
and north-west India at BMNH (DP). In size and structure the type series clearly concurs
with the Mongolian race.
Discussion
There is no reason to suppose that the isabellinus type series, collected on the wintering
grounds, is from a hybrid population. The only question concerns which of two sandy
isabelline races it represents, speculigerus from Mongolia or the pale-primaried race breeding
in the Tarim Basin, Xinjiang. (We can reasonably discount tsaidamensis from further east in
China from the argument.) The dark brown to blackish primaries of all three Berlin males
place them firmly with Mongolian birds, as does the large speculum of paralectotype ZMB
1889, and the complete black loral band of lectotype ZMB 1887 and paralectotype ZMB
1889. The loral / forehead pattern and speculum size of the lectotype, and the incomplete
mask of paralectotype ZMB 1890, may not be typical of speculigerus but nonetheless fall
within the normal range of variation associated with this form. The longer wing and lower
tail / wing ratio of the type specimens also accord with the Mongolian rather than the
Tarim Basin race. Thus, plumage and structure confirm that the former should be named
isabellinus, and arenarius therefore be used for the latter.
Acknowledgements
We are grateful to Robert Prys-Jones and Mark Adams for facilities afforded us at the Natural History
Museum, Tring; other museums have been equally helpful, notably the American Museum of Natural
History (AMNH), New York (Joel Cracraft, Paul Sweet), Musee Nationale d'Histoire Naturelle (MNHN),
Paris (Claire Voisin), Zoological Institute, Almaty (Andrey Gavrilov), Zoological Museum, Moscow (Pavel
Tomkovich, Eugeniy Koblik) and Naturhistoriska Riksmuseet, Stockholm (Goran Frisk, Ulf Johansson).
At ZMB we thank Hannelore Lundsberg (ZMB) for providing archive documents and Pascal Eckhoff for
technical assistance in the collection. We are very grateful to Paul Leader and Magnus Hellstrom for kindly
providing photographs.
References:
Cramp, S. & Perrins, C. M. (eds.) 1993. The birds of the Western Palearctic, vol. 7. Oxford Univ. Press.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
David Pearson et al.
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Bull. B.O.C. 2012 132(4)
Hemprich, F. G. & Ehrenberg, C. G. 1833. Fol. E in Ehrenberg, C. G. (ed.) Symbolae physicae seu icones et
descriptiones avium quae ex itinere per Africam Borealem et Asiam Occidentalem. Decas Prima (& pis. Decas
Secunda). Officina Academica, Berlin.
Koblik, E. Av Red' kin, Y. A. & Arkhipov, V. Y. 2006. [Checklist of the birds of the Russian Federation ]. KMK
Scientific Press, Moscow. [In Russian.]
Korelov, M. N. 1970. Family Laniidae. Pp. 364-399 in Dolgushin, I. A. & Korelov, M. N. (eds.) [Birds of
Kazakhstan ], vol. 3. Nauka of the Kazakh SSR, Alma-Ata. [In Russian.]
Kryukov, A. P. 1995. Systematics of small Palearctic shrikes of the 'cristatus group'. Pp. 22-25 in Yosef, R. &
Lohrer, F. E. (eds.) Shrikes (Laniidae) of the world: biology and conservation. Proc. West. Found. Vert.
Zool. 6(1).
Lefranc, N. & Worfolk, T. 1997. Shrikes : a guide to the shrikes of the world. Pica Press, Robertsbridge.
Lichtenstein, [M.] H. [C.] 1854. Nomenclator avium Museu zoologici berolinensis—Namenverzeichniss der in der
zoologischen Sammlung der Koniglichen Universitdt zu Berlin aufgestellten Arten von Vogeln nach den in der
neueren Systematik am meisten zur Geltung gekommenen Namen und Gattungen und ihrer Unterabtheilungen.
Konigliche Akademie der Wissenschaften, Berlin.
Olivier, G. 1944. Monographic des pies-grieches du genre Lanius. Rouen.
Panov, E. N. 1996. Die Wiirger der Palaarktis. Second revised edn. Neue Brehm Biicherei, Westarp
Wissenschaften, Magdeburg.
Panov, E. N. 2009. On the nomenclature of the so-called Isabelline Shrike. Sandgrouse 31: 163-170.
Pearson, D. J. 1979. The races of the Red-tailed Shrike Lanius isabellinus occurring in East Africa. Scopus 3:
74-78.
Pearson, D. J. 2000. The races of the Isabelline Shrike Lanius isabellinus and their nomenclature. Bull. Brit.
Orn. Cl. 120: 22-27.
Stegmann, B. 1930. Uber die Formen der palaarktischen Rotriicken- und Rotschwanzwiirger und deren
taxonomischen Wert. Orn. Monatsb. 38: 106-118.
Stepanyan, L. S. 1990. [Conspectus of the ornithological fauna of the USSR]. Nauka, Moscow. [In Russian.]
Stresemann, E. 1954. Flemprich und Ehrenberg. Reisen zweier naturforschender Freunde im Orient
geschildert in ihren Briefen aus den Jahren 1819-1826. Abh. Dt. Akad. Wiss. Berlin, Klasse Mathem. Allgem.
Naturwiss. 1: 1-177.
Stresemann, E. & Stresemann, V. 1972. Uber die Mauser in der Gruppe Lanius isabellinus. J. Orn. 113: 60-75.
Vaurie, C. 1959. The birds of the Palearctic fauna. Order Passeriformes. H. F. & G. Witherby, London.
Addresses: David Pearson, 4 Lupin Close, Reydon, Southwold, Suffolk, UK, e-mail: djpearson@dsl.pipex.com.
Lars Svensson, S:ta Toras vag 28, S-269 77 Torekov, Sweden, e-mail: lars@lullula.se. Sylke Frahnert,
Museum fur Naturkunde, Leibniz-Institut fiir Evolutions- und Biodiversitatsforschung, Invalidenstrafie
43, D-10115 Berlin, Germany, e-mail: Sylke.Frahnert@mfn-berlin.de
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Andrew B. Black
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Collection localities of the Night Parrot Pezoporus
( Geopsittacus ) occidentalis (Gould, 1861)
by Andrew B. Black
Received 18 May 2012
Summary. — The vanishingly rare Night Parrot Pezoporus occidentalis of the extensive
arid regions of continental Australia is represented by 25 extant specimens in the
world's museums, of 28 known to have been collected. Apart from one collected
before the holotype, the holotype itself, two initially acquired alive and the most
recent two, found desiccated, all were collected during the decade 1871-81 by
Frederick William Andrews. He took almost all of his specimens in the Gawler
Ranges, western South Australia, but one (and possibly more) came from Cooper
Creek in north-eastern South Australia. He alone is known to have actively
searched for Night Parrots and seen the species alive; most, if not all, subsequent
encounters have occurred by chance.
Forshaw et al. (1976) documented then known museum specimens of the Night Parrot
Pezoporus occidentalis (numbered 1-22 in Table 1) and drew attention to the significance of
F. W. Andrews as a collector of up to 16 of them, from the Gawler Ranges and Lake Eyre
regions of South Australia (SA). Two other specimens were prepared following the demise
of birds acquired alive by the Zoological Society of London. The first came from Nonning
Station, Gawler Ranges, in 1867; captured by the pastoralist Charles Ryan it was forwarded
via the distinguished colonial botanist Dr Ferdinand Mueller (Table 1: 15). The other was
acquired in 1873 from an unnamed source via a London dealer (Table 1: 13) (Sclater 1867,
1873, Murie 1868). Among others discussed by Forshaw et al. (1976) a specimen from north-
eastern SA (Table 1: 16), in fact the earliest collected, had been misidentified as the closely
related Eastern Ground Parrot P. wallicus, avoided John Gould's attention and was long
overlooked. The remaining three specimens, including the holotype, were considered to
have come from Western Australia (WA). Forshaw et al. (1976) referred to the Night Parrot
as a 'vanished species', only one specimen having then been collected in the 20th century
(Table 1: 23), near a tributary of the Ashburton River (WA) by Martin Augustus Bourgoin,
but it was poorly prepared and was not retained (Wilson 1937). While there have been no
confirmed sightings of live birds since those of Bourgoin and others named by Wilson (1937)
two carcasses have been found in western Queensland (Table 1: 24-25) (Boles et al. 1994,
Cupitt & Cupitt 2008, McDougall et al. 2009). The archives of the South Australian Museum,
Adelaide (SAMA), contain information (unavailable to Forshaw et al. 1976) concerning the
exchange of Night Parrot specimens and at least one other (in Stuttgart) was overlooked
by them (R. Prys-Jones pers. comm.; Sclater 1894). My aim here is to clarify and extend the
documentation of museum specimens and their localities provided by Forshaw et al. (1976).
Methods
I examined the archives at SAMA for any reference to the acquisition or exchange of
Night Parrot specimens and enquired via the European Curators' Bulletin Board for any
specimens in museum collections that were not listed by Forshaw et al. (1976).
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Bull. B.O.C. 2012 132(4)
TABLE 1
Currently known Night Parrot Pezoporus occidentalis specimens; those that Forshaw et al. (1976) concluded
were probably taken by F. W. Andrews are indicated *.
Museum and registration details Locality Date and comments
Australian Museum, Sydney
1
AM 0.17831
2
AM 0.17832
Gawler Ranges, SA *
12 November 1872 [presumed collection date]
3
AM 8393
'Central SA' *
A mount, registered between 1869 and 1875
4
AM A.9308
5
AM A.9309
Gawler Ranges, SA *
Registered 1880
AM A.9310
Gawler Ranges, SA
Registered 1880; not found in museum (see
below)
Macleay Museum, Sydney
6
B 1618
South Australia *
Sent from SAMA, June 1874
Museum Victoria, Melbourne
7
NMV B36256
Lake Eyre, SA *
Donated by J. Leadbeater, July 1876
8
NMV HLW 54
9
NMV HLW 55
Gawler Ranges, SA *
From the Robert Grant (private) collection
South Australian Museum, Adelaide
10
SAMA B8118
Gawler Ranges, SA *
F. W. Andrews undated
11
SAMA B24172
Gawler Ranges SA *
F. W. Andrews '1880' [A9310 above, returned
from Australian Museum, Sydney, 20 June
1951]
Natural History Museum, Tring
12
BMNH 1868.1.37.30
Mount Farmer, WA
Holotype, collected by C. Farmer, 23
September 1854
13
BMNH 1939.12.9.554
Unknown
Acquired alive by London Zoo, 16 March
1873
14
BMNH 1881.11.7.1117
'Western Australia'
Part of J. T. Cockerell collection
15
BMNH 1868.4.15.3
Gawler Ranges, SA
Received by London Zoo, November
1867. Sternum and shoulder bones; mount
subsequently registered as BM 1990.7.1
National Museums Liverpool
16
LIVM D.640c
North of Cooper Creek, SA
J. McDouall Stuart, 15 October 1845
Museum National d'Histoire Naturelle, Paris
17
MNHN CG 1879, 679.
18
MNHN CG 1879, 680
Unknown *
Received from SAMA, 1879
American Museum of Natural History, New York
19
AMNH 623832
'Western Australia'
Formerly BM 1881.11.7.1118, Cockerell
collection
20
AMNH 623833
South Australia *
'About 1890?' 'collected by Andrews between
Lake Acraman and Lake Gairdner'; type of
Geopsittacus occidentalis whiteae Mathews, 1915
National Museum of Natural History, Washington
21 USNM 71792 South Australia * Exhibited at Philadelphia Exhibition 1876
Museum of Comparative Zoology, Cambridge, MA
22 MCZ 31516 South Australia * Bought by R. A. Ward, dealer, in 1880 or
earlier
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Bull. B.O.C. 2012 132(4)
Additional specimens
23
Never accessioned
Nichol Spring, WA
Collected by M. A. Burgoin, 21 September
1912, near Ashburton River, now lost
24
Queensland Museum QM O
29055
North of Boulia, west
Queensland
17 October 1990 (Boles et al. 1994)
25
Queensland Museum QM O
32613
Diamantina National Park,
west Queensland
17 September 2006 (Cupitt & Cupitt 2008)
26
Staatliches Museum fur
Naturkunde, Stuttgart, specimen
Gawler Ranges?
Probably acquired in 1883 (see text)
27
Strasbourg MZS 14583
Gawler Ranges, SA
Acquired late 19th century (see text)
28
Hungarian Natural History
Museum, Budapest, specimen or
specimens
Unknown
Purchased by G. A. Frank from a London
trader in 1882
Results
SAMA records reveal that collections of birds and mammal from the Gawler Ranges
were acquired from Andrews on up to eight occasions in 1870-83. He sold his first Night
Parrot specimen to the museum in December 1871. Other documented acquisitions were
two in March 1873, six in October 1873 and four in November 1873. The last ten were
listed for possible exchange with the Calcutta museum, but I cannot confirm that any were
used for such a purpose. Night Parrots are not listed with Gawler Ranges collections from
Andrews in 1878, 1880 and 1881, but it is probable that he was successful on at least some
trips in that period because, during September-December 1883, he made several references
(in extant letters) to his continued anticipation of obtaining the species; however, it is
evident that his expectation was unfulfilled in that year.
Andrews supplied SAMA with a single Night Parrot specimen from Cooper Creek
during his engagement as naturalist with the 1874-75 survey of Lake Eyre (SA) and this was
exchanged with John Leadbeater of Melbourne in January 1876 (now in Museum Victoria,
Table 1: 7) as reported by Forshaw et al. (1976). There are records of only four specimens
Figure 1. Strasbourg Night Parrot Pezoporus occidentalis specimen (MZS 14583), for which the English-
language label reads: 'Sp 442 Gould. Geopsittacus occidentalis. Gawler Ranges S. Aust.' (© M. Meister)
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Bull. B.O.C. 2012 132(4)
being exchanged with other collections; one went to the Macleay Museum, University
of Sydney, in June 1874 (Table 1: 6), one to Leadbeater (above) and two from the Gawler
Ranges were forwarded to J. T. Cockerell of Brisbane in September 1873. Forshaw et al.
(1976) found that a registered specimen, listed in the Australian Museum, Sydney (AM
A.9310) was missing but noted a second specimen in SAMA that was unreported by Sutton
(in Wilson 1937). The AM skin had been transferred to SAMA in 1951 (Table 1: 11).
Responses from the European Curators' Bulletin Board enquiry revealed the existence
or former existence of at least three Night Parrot specimens additional to those reported by
Forshaw et al. (1976).
Confirming Sclater's (1894) reference to one or more specimens in Stuttgart, the
museum's catalogue lists one Night Parrot specimen acquired with a grasswren (Amytornis)
specimen from the Gawler Ranges (Black unpubl.). However, it is now missing, presumed
stolen with other parrot specimens (F. Woog in litt. 2012). Documentation of the grasswren
skin indicated its collection in South Australia and purchase in 1883 from Gustav Schneider,
trader and former museum curator in Basel (K. Schwarz pers. comm.). SAMA records show
that its Director Wilhelm Haacke forwarded 1,000 zoological specimens to Schneider in
1882-84 in exchange for specimens from Africa and South America (Hale 1956).
In the Museum of Zoology, Strasbourg, is a well-preserved Night Parrot skin (MZS
14583; Table 1: 23; Fig. 1) whose origin is the Gawler Ranges, SA; the specimen is undated,
but its Strasbourg label is typical of those used in the second half of the 19th century (M.
Meister in litt. 2012).
Records in the Hungarian Natural History Museum, Budapest, show that one or two
Night Parrot specimens were purchased by G. A. Frank from a London-based trader in
1882. However, this collection was destroyed by fire caused by Russian artillery in 1956 (T.
Fuisz in litt. 2012).
In the Department of Zoology, Cambridge University, there is a partial pectoral
girdle and sternum of a Night Parrot (among Newton's sterna collection; UMZC 1007.a,
28.11.1901: M. Brooke pers. comm.). Such skeletal remnants could be from a specimen
already accounted for and are therefore not included in the Table.
Discussion
Frederick William Andrews was a self-employed naturalist who collected (chiefly)
vertebrate specimens for SAMA between 1864 and 1884. He is recognised, in addition to
his unparalleled success with the Night Parrot, as the collector of the syntypes of Eyrean
Grasswren Amytornis goyderi (Gould, 1875).
In their review of Night Parrot specimens Forshaw et al. (1976) identified 23, including
one that was subsequently lost; they emphasised Andrews' success in collecting 16 of the
total. I find that, in addition to two specimens obtained more recently, at least three others
entered European collections during the 19th century; I can also clarify some uncertainties
presented in the earlier review. Of the 19 identified as taken in SA, Forshaw et al. (1976)
showed that one (Table 1: 16), taken by John McDouall Stuart in 1845 during Charles Sturt's
inland expedition, was collected prior to the holotype. Two (Table 1: 13, 15) were sent alive
to London Zoo, one in 1867 from the Gawler Ranges and the other in 1873 from an unknown
locality. The remaining 16 were regarded as being collected by Andrews, including the two
in Paris (Table 1: 17-18), which formed part of a consignment of natural history material
sent from SAMA to the Paris International Exhibition of 1878 (pers. data).
Apart from the holotype, two possible WA specimens were identified by Forshaw et al.
(1976): one in the Natural History Museum, Tring (Table 1: 14) and another in the American
Museum of Natural History, New York (Table 1: 19). Both were presented to the British
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Andrew B. Black
281
Bull. B.O.C. 2012 132(4)
Museum in 1881 by F. D. Godman from the collection of John T. Cockerell (Sharpe 1906).
While Cockerell is not known to have collected in WA, his son James F. Cockerell did so, for
Samuel White in 1879, but with no report of a Night Parrot (Whittell 1954; SAMA archive). J.
T. Cockerell's labels have been found to be inadequate or inaccurate (Sharpe 1906, Whittell
1954; pers. obs.) and, since he held two Gawler Ranges specimens, obtained from SAMA in
September 1873, it is probable that these were the two acquired by Godman.
Two Tring specimens are of birds obtained alive by the Zoological Society of London.
The first (Table 1: 15) was captured in the Gawler Ranges, sent to the Society by Dr Mueller
in 1867 and was well studied in life (Sclater 1867, Murie 1868). The second 'addition to
the menagerie' was obtained alive by the Society from a dealer on 16 March 1873 (Sclater
1873) but died a few days later according to its label (Table 1: 13). It was acquired by Lord
Rothschild and passed to the museum as part of his bequest in 1937 (R. Prys-Jones in litt.
2012). Olsen (2009) believed that the latter was probably obtained by Andrews and, while I
can find no evidence that he took or traded live birds Andrews (1883) did report that 'one
of these parrots was caught in a hut' and was kept in 'a box with a handful of dry grass' in
which it subsequently 'concealed itself.
Of the three previously unknown specimens in European collections reported here,
only one can now be traced. The missing Stuttgart specimen was acquired in 1883, with
another bird specimen from the Gawler Ranges. The documentation associated with that
in Strasbourg indicates that it too was from the Gawler Ranges. The Budapest specimen(s)
are missing, but records there provide a year (1882) consistent with the period in which
Andrews' specimens were arriving in Europe.
Adding the two Cockerell skins and three European specimens to Forshaw et al.'s (1976)
total of 16 indicates that Andrews was very probably responsible for 21 of the 28 known
Night Parrot specimens, and there may well have been more, such as the second London
Zoo bird. Yet SAMA records list only 13 acquired by the museum and only four exchanged
with other collections (Table 1: 6-7, 14, 19). Almost certainly the records are incomplete, but
an additional explanation exists for which Forshaw et al. (1976) provided some evidence.
Three of the specimens in the Australian Museum, Sydney, were sold to its Curator, E. P.
Ramsay by SAMA Curator, F. G. Waterhouse, not as an exchange between museums. In his
monthly reports to the museum's board Waterhouse made frequent reference to the lack of
adequate finances for purchases, even once stating that he had used personal funds for the
purpose. It seems likely that he also made direct sales to raise money, but such transactions
do not appear in his monthly reports or in the museum's annual reports (pers. data). It is
possible that the two specimens in the H. L. White collection, Museum Victoria, acquired
from the Robert Grant collection, presumably in 1894 (Forshaw et al. 1976), were obtained
similarly, and the Harvard and Budapest specimens were certainly purchased via dealers.
Another possibility is that Andrews himself might have sold specimens to individuals or
to dealers; he was self-employed and is known to have made private sales (Hale 1956). In
his one publication concerning Night Parrots, Andrews (1883) stated that he had 'shot some
specimens at Cooper's Creek in 1875' yet only one was acquired by SAMA. In view of this
uncertainty, while the great majority of his specimens were from the Gawler Ranges, one
or more that lack locality details could be from Cooper Creek.
Acknowledgements
I thank Robert Prys-Jones for suggesting and then assisting my enquiry of European museum collections,
and all of the collection managers who responded, including those named herein who provided positive
information. I am grateful to Wayne Longmore and an anonymous reviewer for constructive suggestions on
the submitted draft.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Andrew B. Black
282
Bull. B.O.C. 2012 132(4)
References:
Andrews, F. W. 1883. Notes on the Night Parrot. Trans. Roy. Soc. South Austr. 6: 29-30.
Boles, W. E., Longmore, N. W. & Thompson, M. C. 1994. A recent specimen of the Night Parrot. Emu 94:
37-40.
Cupitt, R. & Cupitt, S. 2008. Another recent specimen of the Night Parrot Pezoporus occidentalis from western
Queensland. Austr. Field Orn. 25: 69-75.
Forshaw, J. M., Fullagar, P. J. & Harris, J. I. 1976. Specimens of the Night Parrot in museums throughout the
world. Emu 76: 120-126.
Hale, H. M. 1956. The first hundred years of the South Australian Museum. Rec. South Austr. Mus. 12: 1-225.
McDougall, A., Porter, G., Mostert, M., Cupitt, R., Cupitt, S., Joseph, L., Murphy, S., Janetzki, H., Gallagher,
A. & Burbidge, A. 2009. Another piece in an Australian ornithological puzzle - a second Night Parrot is
found dead in Queensland. Emu 109: 198-203.
Murie, J. 1868. On the nocturnal ground-parakeet ( Geopsittacus occidentalis Gould). Proc. Zool. Soc. Eond. 1868:
158-165.
Olsen, P. 2009. Night Parrots: fugitives of the inland. Pp. 121-146 in Robin, L., Heinsohn, R. & Joseph, L. (eds.)
Boom and bust: bird stories for a dry country. CSIRO Publishing, Collingwood.
Sclater, P. L. 1867. [Additions to the Society's Menagerie.] Proc. Zool. Soc. Lond. 1867: 890-891.
Sclater, P. L. 1873. [Additions to the Society's Menagerie]. Proc. Zool. Soc. Eond. 1873: 433-434.
Sclater, P. L. 1894. Ornithology at Munich, Stuttgart, Darmstadt, Frankfort and Cassel. Ibis (6)6: 106-108.
Sharpe, R. B. 1906. The history of the collections contained in the natural history departments of the British Museum.
London.
Whittell, H. M. 1954. The literature of Australian birds: a history and a bibliography of Australian ornithology.
Paterson Brokensha, Perth.
Wilson, H. 1937. Notes on the Night Parrot, with references to recent occurrences. Emu 37: 79-87.
Address: South Australian Museum, North Terrace, Adelaide 5000 SA, Australia, e-mail: abblack@bigpond.
com
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
283
Bull. B.O.C. 2012 132(4)
Morphological differentiation and speciation among
darters ( Anhinga )
by Richard Schodde, Guy M. Kirzvan & Richard Porter
Received 21 June 2012
Summary.— Morphological analysis of the major populations of Old World darters
(. Anhinga spp.) from Africa to Australia identified significant regional differentiation
in sexual dimorphism, shape and structure of the pale cheek stripe, pattern and
tone of the chin and foreneck, form of scapular feathering, and marking and tone
of the greater wing-coverts. Further differentiation was found in the proportions
of tail, bill and feet, and in bare-part colours, particularly in the irides and feet.
Among the three major forms— Afro-Middle East rufa, Oriental melanogaster and
Australasian novaehollandiae— qualitative differentiation in plumage patterning
was almost of the same high order as that between these darters and the Anhinga
A. anhinga of the New World. Furthermore, differentiation of a lower but still
clear-cut order was found among African, Malagasy and Middle East populations
of rufa, and between Australian and New Guinean populations of novaehollandiae.
We conclude that rufa, melanogaster and novaehollandiae have speciated, and that
Malagasy, Middle East and New Guinean isolates are subspecies of African rufa
and Australian novaehollandiae respectively.
It is conventional today to recognise two species of Anhinga : the Anhinga A. anhinga in
tropical and subtropical America and the Darter A. melanogaster from Africa across southern
Asia to Australasia in the Old World (Voous 1973, Wolters 1975, Cramp & Simmons 1977,
Dorst & Mougin 1979, Brown et al. 1982, M. D. Bruce in White & Bruce 1986, Marchant &
Higgins 1990, Orta 1992, Johnsgard 1993, Inskipp et al. 1996, Wells 1999, Dickinson 2003).
Yet while the status of the American anhinga has hardly been in question, the populations
of Old World darters have been in a state of taxonomic flux throughout the later 20th
century. Peters (1931), first to employ the polytypic species concept in a global list of birds,
treated them as three species: Afro-Middle East rufa Daudin, 1802, Oriental melanogaster
Pennant, 1769, and Australian novaehollandiae Gould, 1847. In rufa, furthermore, he
distinguished three subspecies: nominate rufa through sub-Saharan Africa, vulsini Bangs,
1918, in Madagascar and chantrei Oustalet, 1882, in the Middle East. Vaurie (1965) and
Mayr & Short (1970) accepted Peters' species, the former concluding that morphological
differentiation among rufa, melanogaster and novaehollandiae was too great for any lumping.
Since then, nevertheless, there has been a cascade of lumping, sparked by Voous (1973),
Wolters (1975) and Condon (1975) placing all Old World forms in one species. They were
promptly supported by Harrison (1978) with his finding of common structure and tendinal
canals in tarsometatarsi, although his sample was small and possibly affected by age.
Except the AOU (1983), Sibley & Monroe (1990), Andrew (1992), Rasmussen & Anderton
(2005) and Kirwan et al. (2008) who maintained the three species of Peters (1931), other
major revisers followed the lead of Voous, Wolters, Condon and Harrison, and demoted
Peters' species to subspecies— see references above. Consequent casualties were most of
Peters' (1931) subspecies, and at times even certain of his species names. Condon (1975)
and Dorst and Mougin (1979), for example, combined African and Australian darters
(rufa, novaehollandiae) in one subspecies separate from the intervening Oriental darter
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
284
Bull. B.O.C. 2012 132(4)
(melanogaster). Zoogeographically it was absurd, justified by the dubious observation that
'Australian birds were virtually indistinguishable from subspecies rufa of Africa' (G. F. van
Tets in Condon 1975; H. T. Condon pers. comm.). Most reviewers (including Kirwan et
al. 2008) also dropped Malagasy and Middle Eastern forms of rufa—vulsini and chantrei—
into synonymy, along with subsequently described papua Rand, 1938, from New Guinea.
Wolters (1975) recognised chantrei, and Orta (1992) and Dickinson (2003) listed vulsini, but
that was all.
Molecular studies have so far added little. MtDNA sequences of African rufa and
Australian novaehollandiae were compared incidentally by Kennedy et al. (2005) in a
methodological study focused on resolving phylogenetic signal. It showed divergence
comparable to levels of mtDNA distance between species of cormorants ( Phalacrocorax )
and boobies ( Sula ). Intervening Oriental melanogaster was not included in the study, yet it
was enough for Christidis and Boles (2008) to treat all three major Old World forms— rufa,
melanogaster and novaehollandiae— as separate species. Apart from the work of Harrison
(1978), which is limited in any case, none of the above reviews is supported by an analysis
of characters across taxa— morphological or molecular. There have, in fact, been no decent
comparative descriptions of how the major continental forms of Old World darters differ
from one another since Ogilvie-Grant's (1898) account over 100 years ago. Towards filling
this gap and providing an explicit rationale for species-group taxonomy in the Old World
darters, we offer here a detailed morphological analysis of all regional forms, together with
conclusions concerning taxonomic status.
Materials and Methods
This study is based on dry skin material of all Old World populations of darters
preserved in the globally rich and comprehensive collections of the American Museum
of Natural History, New York (AMNH), augmented by series' in Australian national and
provincial museums: Australian Museum, Sydney (AM); Australian National Wildlife
Collection, Canberra (ANWC); Museum Victoria, Melbourne (MV); Queensland Museum,
Brisbane (QM); South Australian Museum, Adelaide (SAMA); and Western Australian
Museum, Perth (WAM). We also examined the small collections of New Guinean material
in the Papua New Guinea National Museum, Port Moresby (PNGNM). These series were
compared among themselves and with a selection of 15 males and 15 females of nominate
A. anhinga in AMNH, as an out-group. Altogether 275 adult specimens of both sexes were
compared. The numbers of each sex of each taxon examined are given in the header to
Table 1, and the numbers of selected specimens measured are given against taxon and sex
in Table 3.
We analysed plumage patterning from long series laid out by region. As well as taking
standard measurements (wing, tail, bill and tarsus) to assess gross size, moreover, we also
calculated ratios to compare allometric proportions, which are usually more significant
indicators of adaptation and differentiation. Wing was measured as flattened chord, tail as
the length of the central rectrices, bill from the tip to feathers on culmen, and tarsus from the
notch on the heel to the top of the knuckle bridging the base of the toes. Bare-part colours,
recorded inconsistently in museum collections, were augmented and teased out for age and
sex from details in handbooks (Palmer 1962, Cramp & Simmons 1977, Brown et al. 1982,
Marchant & Higgins 1990) and photographs (Orta 1992; internet, only for taxa identifiable
by plumage).
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
285
Bull. B.O.C. 2012 132(4)
Morphological analysis
Qualitative differentiation in plumage patterns.— The results of plumage analysis are
detailed in Table 1. They show a pattern of regional differentiation in principally three
sets of characters: (1) face and throat pattern, (2) upperwing pattern and (3) gross ventral
pattern. Differing from plain-headed American anhinga, all Old World populations share a
pale cheek-side neck stripe in both sexes in both breeding and non-breeding plumages. But
there the similarity among them ends. In rufa, vulsini and chantrei, the stripe is moderately
long and slender, in melanogaster still longer and narrower, but in novaehollandiae and papua
short and broad. Furthermore, the structure of its feathering differs. In melanogaster and
the Australasian group, its pennae resemble feathering elsewhere on the face and neck in
all seasons, but in rufa, vulsini and chantrei, they become distinctively if shortly plumose
during breeding (Cramp & Simmons 1977) and terminate in short rust-brown hairs as they
wear. Such pluming may be homologous with the longer white, rufous and black plumes
that develop on the sides and back of the head and neck in breeding male and female
anhinga. On the throat, breeding males of all regional forms differ as detailed in Table 1,
with primary differentiation between rufa / vulsini (rufous with ill-defined white border
to chin), melanogaster (spotted blackish on white), novaehollandiae / papua (black with clear
broad white border) and anhinga (plain black). The creamy throat of male chantrei may
reflect more the retention of non-breeding foreneck tone than any substantive difference in
pattern from the rufa group.
In upperwing pattern, anhinga again stands out in the brilliance of its broad silver-grey
sash across the shoulders (wing-coverts). The effect of this grey sash, centred on the greater
wing-coverts, is enhanced by a lack of black bordering to the feathers there. It produces a
contrasting three-band wing pattern: a distal all-black remige band and a proximal inner
covert band of black spotted silver, split by a broad plain silver median band through the
greater coverts. This sash is present in all Old World regional groups, but is never as broad
and bright as in anhinga, and its feathering is bordered with black on the greater as well as
the inner coverts (Table 1). In novaehollandiae and papua it is still distinct, and indeed paler,
broader and rather anhinga- like in females, but because of black feather edging and darker
toning, the greater wing-coverts in males blend with the inner and together contrast with
the all-black remiges in more of a two-band pattern. There is no sexual dimorphism in
melanogaster, but the grey tone to the feather centres is as silvery as in female novaehollandiae
/ papua or even whiter, producing a more contrasting two-band pattern against the all-black
body of both sexes. In both sexes of rufa, and to a lesser extent of vulsini and chantrei, the pale
greater wing-coverts band is narrowest and dullest of all, even fuscous in tone. It neither
stands out as a discrete band as in anhinga and female novaehollandiae / papua, nor blends with
the inner coverts to contrast with the remiges as in melanogaster and male novaehollandiae /
papua. Rather, it has the appearance of a dull narrow divider between the inner coverts and
remiges in an almost monotone wing pattern. Differences in the upperwings extend to the
scapulars which are very long and attenuate with dull buffy-grey shafts in rufa and vulsini,
similar in form but with silvery shafts in chantrei and melanogaster, rather short, abruptly
acute and dull grey-shafted in Australasian novaehollandiae and papua, and of intermediate
form and silvery shafted in anhinga.
Seasonal cycles, sexual dimorphism and ontogeny complicate the expression of gross
ventral pattern. It is simplest in adults of the Afro-Oriental forms, in which non-breeding
plumage resembles breeding, and females resemble males except for paler and duller toning
respectively (Cramp & Simmons 1977). In these populations, black over the lower ventral
region extends to the top of the lower foreneck, where it is abruptly demarcated from
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
TABLE 1
Traits of adult breeding plumage in Old World regional forms of Anhinga, and of American A. anhinga.
Most but not all adults in the sample sizes quoted for each form are in breeding plumage.
Richard Schodde et al.
286
Bull. B.O.C. 2012 132(4)
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© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Pale tail tip absent absent absent absent absent absent present
Richard Schodde et al.
288
Bull. B.O.C. 2012 132(4)
variously rufous, brown and cream tones over the rest of the foreneck. American anhinga,
in contrast, is markedly sexually dimorphic. Adult males are all black in both breeding
and non-breeding plumages, whereas females resemble Afro- Asian males and females in
pattern and seasonal tones except for a much lower placed demarcation between the tawny-
rufous foreneck and black lower underparts across the mid breast. Sexual dimorphism
and seasonal variation is perhaps most complex of all in Australasian novaehollandiae and
papua. In this group, adult males are entirely black when breeding, except for a small patch
of rich chestnut confined to the mid foreneck. At other seasons, they remain black to the
top of the lower neck, indicative of affinity with Afro-Oriental forms, but become pale to
dull deep rufous over the entire mid and upper foreneck. Adult females, by contrast, are
entirely greyish dorsally and whitish ventrally, and differ only with season by developing
clear black margins to the cheek stripe when breeding. Marchant & Higgins (1990) did not
recognise breeding and non-breeding plumages in novaehollandiae , but they are identified
here from correlating three-year-old birds or older (fully crimped central rectrices and
scapulars) with their gonad condition in material in ANWC.
American anhingas have been thought to differ from all Old World forms in their
narrowly whitish- tipped tails (Ogilvie-Grant 1898). It is noteworthy then that juveniles of A.
rufa have been described (Cramp & Simmons 1977) and figured (Cramp & Simmons 1977,
pi. 26, Brown et al. 1982, pi. 8) with pale tips to the tail. Juveniles of Australian (n = 10) and
New Guinean ( n = 3) populations lack this bar (Marchant & Higgins 1990: 827). Thus a pale
terminal tail bar is evidently present in juveniles of some Old World populations, but not
others, and is lost in all with age. Such changes occur throughout the plumages of the genus.
Juveniles of all Old and New World forms are uniformly pale whitish ventrally and dull
grey dorsally, and appear to gain adult and sexual plumages progressively into their late
second and third years (Palmer 1962, Marchant & Higgins 1990). Different age classes from
different regions overlap in these traits during development, because of which analyses
here are based on adults with fully crimped central rectrices and scapulars, and particularly
those in breeding plumage.
Variation in hare-part colours.— Data for bare-part colours in adults of all regional
populations (Table 2) reveal significant inter- taxon differentiation in colour of the irides,
gular skin and feet, and minor differences in colour of bill and face. Some but not all
differentiation is linked with breeding and sex. In the irides, colour is consistently yellow
in melanogaster, novaehollandiae and papua, and deep red in anhinga at all times, whereas it
apparently varies from yellow to red in Afro-Middle East forms (Cramp & Simmons 1977).
Bill colour varies little with age, sex and season among regional forms, although it is richest
in breeding adults and darker over the maxilla in males, particularly in breeding anhinga ,
enhancing the yellow of the mandible in an otherwise blackish head. Face skin tone is a
dull pale yellowish in non-breeders of all taxa except anhinga but deepens and brightens
in Old World populations during breeding, becoming dark yellow particularly in males.
In anhinga, the face of non-breeders is variously dull grey, but becomes rich emerald to
turquoise in breeding males, less so in females. Gular skin, basically yellow, also varies,
becoming blackish in the centre and base in breeding anhinga and breeding Afro-Oriental
forms, although the tone may be more localised and temporary in the latter. In Australasian
populations, however, it appears to remain wholly yellowish at all times. Feet differ among
regions as well. In anhinga they are prevailingly greyish black to black, but in Australasian
populations pallid yellowish flesh at all times. Feet colours in Afro-Oriental forms are
yellowish, approaching Australasian forms, but are washed consistently greyer and are
usually darker. Juveniles of all forms are dull and nondescript in all bare-part colours.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists7 Club
Richard Schodde et al.
289
Bull. B.O.C. 2012 132(4)
TABLE 2
Soft-parts colours of adult Old World regional forms of Anhinga and of American A. anhinga. Colours
recorded are averaged for easier comprehension. Sexes are separated only where colours between them
differ significantly. As a rule, immatures have the colours of non-breeding females and are dull. For
sources, see Materials and Methods.
Taxon
Seasonal
condition
Iris
(both sexes)
Bill
(both sexes)
Facial skin
Gular skin
(both sexes)
Tarsus/toes
(both sexes)
rufa
(Sub-Saharan
Africa)
breeding
variable: rich
yellow to red
rich yellow,
maxilla
washed horn
dusky yellow
(both sexes)
(greenish)
black
dusky brown,
webs blackish
non-breeding
as breeding but
duller
paler, duller
yellow
dull creamy
yellow
dull cream in
both sexes,
sometimes
washed black
deep grey-
brown to
yellowish
brown, webs
yellower
vulsini
(Madagascar)
breeding
yellow (? to
red)
rich yellow,
maxilla
washed horn
dull yellow
(both sexes)
blackish
pale grey with
yellow wash,
webs yellower
non-breeding
as breeding but
duller
paler, duller
yellow
as breeding or
duller
dull cream
pale greyish
yellow, webs
yellower
chantrei
(Middle East)
breeding
variable: yellow
to red
rich yellow,
maxilla
washed horn
dull yellow
(both sexes)
blackish
pale greyish
yellow, webs
yellower
non-breeding
as breeding but
duller
paler, duller
yellow
dull cream
dull cream
pale greyish
cream with
flesh wash
melanogaster
(South-East
Asia)
breeding
bright yellow
rich yellow,
maxilla
washed horn
mid to dark
yellow (both
sexes)
rich yellow
with variable
black wash
pale yellow
to dark grey,
webs yellower
non-breeding
dull yellow
paler, duller
yellow
pale creamy
yellow
pale creamy
yellow with
occasional
black wash
pale cream to
grey-yellow,
webs yellower
novaehollandiae
(Australia)
breeding
bright to deep
yellow
rich yellow,
maxilla
washed
greenish horn
dark yellow
in <?<?, bright
yellow in ? ?
mid yellow
pale yellowish
flesh, webs
browner
yellow
non-breeding
dull yellow to
pale brown
paler, duller
yellow
pale greyish
yellow (both
sexes)
pale cream
pale greyish
flesh, webs
yellower
papua
(New Guinea)
breeding
as
novaehollandiae
as
novaehollandiae
as
novaehollandiae
as
novaehollandiae
as
novaehollandiae
non-breeding
as
novaehollandiae
as
novaehollandiae
as
novaehollandiae
as
novaehollandiae
as
novaehollandiae
anhinga
(New World)
breeding
rich deep red
rich yellow,
maxilla
washed
variably
dusky
emerald to
blue around
eye in
duller in ? ?
black
olive-black
non-breeding
duller red
dull
horn-yellow
dull
grey-green
pale greyish
yellow
greyish yellow
in c ?<?, dull
yellow-grey
in ??
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
290
Bull. B.O.C. 2012 132(4)
which are generally of the same tone in non-breeding females. As a rule, the bare parts in
breeding males are deeper and more intensely toned than in females.
Morphometric differentiation in size and proportions.— Body size, as estimated by wing
length (Table 3), differs little among regional Old World forms and nominate American
anhinga. Furthermore, there is little sexual dimorphism in most forms, particularly the
Australasian group. It is most marked in American anhinga , in which males are larger
than females, and in Malagasy vulsini, in which females are larger than males in all parts.
Measurements for vulsini may be biased by small sample size and the immaturity of two
of the four males, although the bills of the latter, always shorter in juveniles, are as long
as those of the two adult males. There are, however, regional differences in gross and
proportional tail, bill and tarsus length. Both American anhinga and Malagasy vulsini are
longest in tail and bill, and shortest in tarsus, all sexually reversed between the two forms.
Sexual dimorphism in proportional bill length is particularly marked in anhinga, suggesting
partitioning of prey. In contrast, Australasian novaehollandiae and papua are shortest in tail
and bill and longest in tarsi, with virtually no sexual dimorphism. The remaining Afro-
Oriental forms, rufa, chantrei and melanogaster, are intermediate in all gross measurements
and proportions, with males averaging slightly larger than females.
Taxonomic synthesis
None of the major regional forms of Old World darters meet, meaning that their
interaction cannot be assessed under the Biological Species Concept. Middle Eastern chantrei
extends east to the lower Tigris-Euphrates wetlands in Iraq and Iran (Khaleghizadeh et al.
2011; RP pers. data, including photographs), and Oriental melanogaster west to the Indus
Valley, but they are separated by almost 2,000 km of unsuitable habitat along the coast and
hinterland of Iran and west Pakistan. In the Indonesian archipelagos, melanogaster extends
east to Sulawesi as a breeder (White & Bruce 1986), while Australasian novaehollandiae ranges
north-west to Timor-Leste and Roti (Trainor 2005a,b) in the Lesser Sundas, c.600 km south-
east of Sulawesi. On Timor-Leste, where Trainor (2005a) found novaehollandiae breeding,
McKean et al. (1975) had earlier recorded nominate melanogaster, suggesting sympatry. But
as McKean et al. (1975) gave no diagnostic details of the darters they saw, identity may
have been presumed and the record requires confirmation (White & Bruce 1986). Elsewhere
in that region only novaehollandiae has been recorded, in both the south Moluccas and
Banda Sea islands, invariably as a vagrant from either Australia or New Guinea (White
& Bruce 1986). New Guinean (papua) and Australian (novaehollandiae) populations of the
Australasian group may mix to some degree in the Trans-Fly region of southern New
Guinea and on the Cape York Peninsula, Queensland. Nevertheless, sampling to date
(Id', 59 9 from the Trans-Fly, 6 dd, 4 9 9 from the Cape York Peninsula) does not show it
morphologically (see below). In the absence of confirmed interaction between contiguous
populations of any taxa, then, estimates of speciation must rely on levels of morphological,
behavioural and molecular differentiation.
The combined morphological data indicate that Old World darters as a group are well
differentiated from American A. anhinga. All lack breeding plumes on the head-sides and
pale-tipped tails when adult, and they share pale cheek stripes, black feather borders to a
narrower pale sash across the greater wing-coverts, and feet and facial colours dominated
by pale flavonoid pigmentation. Levels of sexual dimorphism in size and proportions are
also low. Yet among themselves, Afro-Middle East, Oriental and Australasian groups are
almost as deeply differentiated as any one of them is from anhinga. The Australasian group
is most divergent with a short, broad cheek stripe, black-necked breeding plumage, with a
broad white chin fringe in males, short scapulars, uniquely white-ventered females in all
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
291
Bull. B.O.C. 2012 132(4)
TABLE 3
Measurements of adults and near adults of all regional forms of Old World Anhinga, and of American
A. anhinga; range plus means (in parentheses). For source and selection of samples, see Materials and
Methods.
Taxon
Sex
n
Wing
Tail
Culmen
Tarsus
Tail/wing
Culmen/
Tarsus/wing
(mm)
(mm)
(mm)
(mm)
ratio
wing ratio
ratio
rufa
dd
18
336-366
220-260
73.0-85.0
39.0-47.3
0.63-0.75
0.205-0.240
0.121-0.133
(Sub-Saharan
(349.7)
(236.4)
(80.0)
(44.8)
(0.68)
(0.229)
(0. 126)
Africa)
??
10
333-357
230-246
70.3-80.7
41.4-46.3
0.66-0.71
0.210-0.240
0.115-0.135
(343.6)
(237.1)
(75.9)
(43.2)
(0.69)
(0.221)
(0.126)
vulsini
dd
4
335-347
235-258
75.2-85.2
43.1-44.3
0.70-0.74
0.217-0.246
0.124-0.131
(Madadascar)
(343.0)
(249.8)
(80.1)
(43.9)
(0.73)
(0. 234)
(0.128)
??
5
343-365
261-280
77.0-92.5
40.6-48.2
0.73-0.77
0.220-0.265
0.116-0.137
(354.2)
(269.0)
(84.6)
(44.2)
(0.76)
(0.237)
(0.125)
chantrei
dd
7
337-359
228-237
80.2-88.3
44.2-47.0
0.65-0.70
0.233-0.257
0.124-0.137
(Middle East)
(347.1)
(232.0)
(84.7)
(45.5)
(0.67)
(0.244)
(0.131)
??
6
328-344
208-228
76.0-85.9
43.4^9.6
0.63-0.68
0.222-0.255
0.127-0.144
(336.2)
(219.5)
(81.5)
(45.8)
(0.65)
(0.242)
(0.136)
melanogaster
dd
18
328-364
215-258
77.0-92.0
34.0-47.5
0.63-0.72
0.230-0.260
0.110-0.140
(South-East
(346.3)
(233.5)
(82.8)
(43.3)
(0.67)
(0.240)
(0.128)
Asia)
??
8
340-360
218-245
67.8-79.4
39.8^5.0
0.62-0.72
0.200-0.230
0.115-0.130
(347.0)
(233.9)
(76.2)
(42.6)
(0.67)
(0.220)
(0.123)
novaehollandiae
d'd'
62
331-368
197-240
66.8-80.0
44.5-52.7
0.59-0.66
0.195-0.230
0.128-0.145
(Australia)
(349.6)
(217.0)
(74.9)
(48.0)
(0.62)
(0.215)
(0.137)
??
63
320-370
200-245
65.0-84.0
45.0-55.0
0.59-0.69
0.190-0.235
0.130-0.152
(346.9)
(220.3)
(75.6)
(48.8)
(0.63)
(0.218)
(0.139)
papua
d'd'
3
347-353
224
67.6-75.2
45.6^9.4
0.64-0.65
0.194-0.214
0.131-0.142
(New Guinea)
(349.3)
(224.0)
(72.4)
(47.7)
(0.65)
(0.207)
(0.136)
??
12
334-360
218-234
73.6-86.0
41.7-50.4
0.62-0.68
0.217-0.239
0.125-0.142
(343.8)
(223.3)
(78.2)
(45.6)
(0.65)
(0.227)
(0.134)
anhinga
dd
10
347-368
246-270
85.0- 96.0
39.5- 44.0
0.71-0.75
0.248-0.270
0.114-0.122
(New World)
(354.0)
(259.4)
(91.8)
(41.5)
(0.73)
(0.260)
(0.118)
??
8
327-354
238-269
75.5- 88.5
37.5- 44.0
0.68-0.77
0.230-0.253
0.112-0.125
(342.1)
(251.8)
(82.0)
(41.0)
(0.74)
(0.240)
(0.120)
plumages, short bills and tails, and relatively long, pale flesh-toned feet. Afro-Middle East
and Oriental populations may be more alike in proportions and bare-part colours, attenuate
cheek stripes and scapulars, and sexually similar breeding plumages, but differentiation
is still marked. Both neck colour and throat patterns differ in breeding plumage, and the
cheek stripe feathering in the Afro-Middle East group is shortly plumose, a state missing
from both Oriental and Australasian groups. Upperwing patterns differ too, and from
the Australasian group. In the latter, it is clearly two-banded and sexually dimorphic, the
spotted grey shoulder band brighter in females than males. In the Oriental, it is sexually
monomorphic yet even more brightly and contrastingly two-banded. But in the Afro-Middle
East group, it is a dull in both sexes, the upperwing almost monotone with fuscous-copper
shoulders except, to some extent, in the Middle East form.
Paedomorphism has played a significant role in differentiation, accounting for the
paler head, neck and bare parts in non-breeding males and females of African and Oriental
populations, creamy neck in all plumages in the Middle East, and grey-backed, white-
ventered adult females in Australasia. Patterns on the upperwing and face and throat
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
292
Bull. B.O.C. 2012 132(4)
in breeding plumage are of a different order, and may be implicated deeply in species
recognition in sexual and social display. Focal elements in such behaviour are 'wing waving'
and 'peering' with sideways head twisting on an outstretched neck (Cramp & Simmons
1977, Brown et al. 1982, Marchant & Higgins 1990). Both sets of movements have the effect of
showing off the face and wing markings; and it follows that unfamiliar patterns there may
hinder mating and serve as barriers to reproduction (cf. Price 2008: 273-297). Accordingly,
we have given extra weight to divergence in nuptial face and upperwing pattern among
the regional populations of Old World darters, leading us to separate Australasian, Oriental
and Afro-Middle East groups as species. They are novaehollandiae, melanogaster and rufa
respectively. Depths of mtDNA sequence divergence, as far as they go, are supportive
(Kennedy et al. 2005). As sister species that represent one another in different geographical
regions, these taxa form a superspecies separate from American anhinga ( pace Dorst &
Mougin 1979, Sibley & Monroe 1990). The latter is differentiated at deeper morphological
levels also (Harrison 1978), a position corroborated by mtDNA data (Kennedy et al. 2005).
Within rufa and novaehollandiae there is further regional differentiation in morphology,
but at a lower level. New Guinean papua resembles novaehollandiae in all plumage patterns,
bare-part colours, size and proportions, but its females are consistently dark dusky-backed,
in contrast to the mid to deep brownish grey tones in Australian females, and the pale shafts
on the scapulars and rich grey centres to the wing-coverts are often reduced. Although such
melanism may have been driven by a more humid environment (Gloger's ecogeographical
rule), the difference is consistent and appears genetically entrenched; thus we agree with
Rand (1938) in recognising papua subspecifically in the novaehollandiae complex. Within the
rufa complex, Malagasy vulsini, though like African rufa in pattern and tone, has a browner
head and rather washed-out cheek stripe with little blackish bordering, particularly in
males, as well as paler, greyer, greater upperwing-coverts. Males also appear to be smaller
than females (Table 3). Although these states may be affected by immaturity in the male
sample screened (crimping on central rectrices is reduced), they are consistent across all
parameters. Middle Eastern chantrei, which, as Kir wan et al. (2008) correctly point out, was
referred in error to A. melanogaster by Sibley & Monroe (1990), appears to be part of the rufa
complex too: it has the distinctive rufa pluming of the cheek stripe, and is rufa-Wke in size,
proportions and sexual dimorphism, with ventral black from the lower neck down in both
sexes. Its neck is nevertheless cream-toned in all plumages— even quite white in males,
against which the black gular line above the cheek stripe stands out. The pale centres to the
upperwing-coverts and scapulars are lighter, more silvery grey as well. Cramp & Simmons
(1977) and Kirwan et al. (2008) included chantrei in rufa because of overlapping variation in
foreneck and wing-coverts tone supposedly reported by Ticehurst, but we cannot confirm
this observation nor, indeed, its reference. Accordingly, we treat vulsini and chantrei as
subspecies of rufa. A. r. chantrei appears to survive today only in the Huweizah (Hawizhe)
and Hoor-al-Azim wetlands within the lower Tigris-Euphrates basin in Iraq and Iran,
although there is one recent record from northern Israel (Ottens 2006) where the species
wintered regularly until 1957 (Shirihai 1996). Its decline and rarity needs recording on
international registers of threatened fauna.
Conspectus of Old World taxa of Anhinga
To conclude, we summarise the taxonomic findings of this study in the following
sequential conspectus of the species and subspecies of Old World darters, together with a
summary distribution of the taxa and suggested English names for the species. All species
are considered to form a superspecies separate from A. anhinga , as indicated by the senior
species-group name in square brackets.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Richard Schodde et al.
293
Bull. B.O.C. 2012 132(4)
Anhinga [ melanogaster ] rufa (Daudin, 1802)— African Darter
A. r. rufa— sub-Saharan Africa
A. r. vulsini Bangs, 1918— Madagascar
A. r. chantrei (Oustalet, 1882)— lower Tigris-Euphrates wetlands (Iraq, Iran), formerly
Lake Antioch = Amik Gold (Turkey) before its draining, and northern Israel (in winter)
Anhinga [ melanogaster ] melanogaster Pennant, 1769 — Oriental Darter
Pakistan (Indus Valley), India, Sri Lanka, mainland South-East Asia (except Malay
Peninsula), Greater Sundas, Philippines, Sulawesi, Sumbawa and ?Timor
Anhinga [ melanogaster ] novaehollandiae (Gould, 1847) — Australasian Darter
A. n. papua Rand, 1938— lowland New Guinea and satellite islands
A. n. novaehollandiae— mainland Australia except central deserts; the subspecific
identity of populations on Timor and Roti (Lesser Sundas , Banda Arc and Moluccas)
remains to be established
Acknowledgements
We record our gratitude to Joel Cracraft and Paul Sweet for facilitating our review of specimen material in the
American Museum of Natural History, New York, and to the curators and collection managers of Australian
and Papua New Guinean national and provincial museums for similar help. T. P. Inskipp, N. W. Longmore
and D. R. Wells provided helpful reviews of the submitted manuscript. The senior author's specimen studies
were supported by Walter Bock, Wayne Longmore and a Collections Grant from the American Museum of
Natural History (Ornithology), New York.
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Peters, J. L. 1931. Check-list of birds of the world, vol. 1. Harvard Univ. Press, Cambridge, MA.
Price, T. 2008. Speciation in birds. Roberts & Co., Greenwood Village, CO.
Rand, A. L. 1938. Results of the Archbold Expeditions no. 19. On some non-passerine New Guinea birds.
Amer. Mus. Novit. 990: 1-15.
Rasmussen, P. C. & Anderton, J. C. 2005. Birds of south Asia: the Ripley guide. Smithsonian Institution,
Washington DC & Lynx Edicions, Barcelona.
Shirihai, H. 1996. The birds of Israel. Academic Press, London.
Sibley, C. G. & Monroe, B. L. 1990. Distribution and taxonomy of birds of the world. Yale Univ. Press, New
Haven, CT.
Trainor, C. R. 2005a. Waterbirds and coastal seabirds of Timor-Leste (East Timor): status and distribution from
surveys in August 2002-December 2004. Forktail 21: 61-78.
Trainor, C. R. 2005b. Birds of Tapuafu Peninsula, Roti Island, Lesser Sundas, Indonesia. Forktail 21: 121-131.
Vaurie, C. 1965. The birds of the Falearctic fauna. Non-passeriformes. H. F. & G. Witherby, London.
Voous, K. H. 1973. List of recent Holarctic bird species. Ibis 115: 612-638.
Wells, D. R. 1999. The birds of the Thai-Malay Peninsula, vol. 1. Academic Press, London.
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Indonesia): an annotated check-list. BOU Check-list No. 7. British Ornithologists' Union, London.
Wolters, H. E. 1975. Die Vogelarten der Erde, Bd. 1. Paul Parey, Hamburg.
Addresses: Richard Schodde, Australian National Wildlife Collection, CSIRO Ecosystem Sciences, G.P.O.
Box 1700, Canberra, ACT, Australia 2601 and Australian Biological Resources Study, G.P.O. Box 787,
Canberra City, ACT, Australia 2601. Guy M. Kirwan, Research Associate, Field Museum of Natural
History, 1400 South Lakeshore Drive, Chicago, IL 60605, USA. Richard Porter, c/o BirdLife International,
Wellbrook Court, Girton Road, Cambridge CB3 ON A, UK.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
William S. Clark
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The eagle genus Hieraaetus is distinct from Aquila, with
comments on the name Ayres' Eagle
by William S. Clark
Received 16 April 2012
Consensus has long existed among taxonomists as to the placement of the species of
the eagles in the genera Aquila and Hieraaetus , the only exception being Wahlberg's Eagle,
which has been treated either as A. wahlbergi or H. zvahlbergi. However, recent comparisons
of DNA sequences of these eagles revealed that both genera were polyphyletic (Helbig et al.
2005, Lerner & Mindell 2005, Griffiths et al. 2007, Haring et al. 2007). To ensure monophyly,
these authors recommended that some species be moved into and out of both genera and
placed Wahlberg's Eagle definitely in Hieraaetus. On the other hand, Sangster et al. (2005)
recommended that Hieraaetus be subsumed into Aquila , for which they cite eight recent
phylogenetic studies, only four of them published in peer-reviewed journals (Roulin
& Wink 2004, Bunce et al. 2005, Helbig et al. 2005, Lerner & Mindell 2005); all of which,
nevertheless, retained Hieraaetus. Thus, none of the four peer-reviewed journal articles
cited by Sangster et al. (2005) to justify placing Hieraaetus into Aquila actually recommended
this. Further, Sangster et al. (2005) proposed only that Booted Eagle ( pennatus ) be included
in Aquila, because their recommendations only considered European birds. The generic
treatment of Sangster et al. (2005) was followed without comment by Gjershaug et al. (2009)
in describing Weiske's or Pygmy Eagle ( weiskei ) as a species separate from Little Eagle H.
morphnoides. Hockey et al. (2005) cited a non-peer-reviewed conference presentation. Wink
& Sauer-Gurth (2000), for subsuming Hieraaetus into Aquila, but their conclusions were
based on the original paraphyletic species.
Wells & Inskipp (2012) advocated moving the three spotted eagles (taxa clanga,
pomarina and hastata) from Aquila into a new genus Aquiloides. They considered that the
genus Aquila that contained the spotted eagles and the taxa formerly in Hieraaetus was
too large, too unwieldy and too diverse. They argued that there are three distinct clades,
Aquila, Hieraaetus and Aquiloides, and that these should be considered separate genera,
with the genera Lophaetus and Ictinaetus in the same clade as the spotted eagles but treated
as monotypic genera because of their divergences from the spotted eagles. Gregory &
Dickinson (2012) reported that the genus name that should be used for the spotted eagles
is Clanga, based on priority.
Brown & Amadon (1968) wrote 'Some recent authors have combined Hieraaetus with
Aquila, but it seems best to keep it separate on the following ensemble of characters, which
admittedly do not hold for every species: size smaller, form more slender, bill smaller, legs
longer, more slender, emargination on primaries deeper.'
In Table 1, I enumerate those characters that separate the species in the revamped
genera Hieraaetus and Aquila. I include characters not mentioned by Brown & Amadon
(1968), but that are important in distinguishing the members of these genera: (1) immature
plumages almost like those of adults, (2) hunting primarily aerial, (3) lack of pale primary
patch, (4) long narrow tail and (5) polymorphism. I believe that the degree of emargination
of the primaries is not a distinguishing character because there is overlap (pers. obs.).
Some species in the revamped genus possess characters not shared by all species, such as
the white headlights of Booted and Ayres' Eagles and the crests of Wahlberg's and Ayres'
Eagles.
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
William S. Clark
296
Bull. B.O.C. 2012 132(4)
TABLE 1
Differences in characters of eagles in the genera Hieraaetus and Aquila. Species are listed in Table 2.
Character
Hieraaetus
Aquila
Size
Small
Medium to large
Immature plumages
Almost like adult
Differ from adult
Hunting style
Primarily aerial
Still and aerial
Pale primary patch
No
Yes
Long narrow tail
Yes
Long on some, but broad
Colour morphs
Dimorphic or polymorphic
Monomorphic, except rapax
TABLE 2
Recommended classification of the genera Hieraaetus and Aquila. * = Type species of genus.
Steppe Eagle Aquila nipalensis
Wahlberg's Eagle Hieraaetus wahlbergi
Eastern Imperial Eagle A. heliaca
Ayres' Eagle H. ayresii
Spanish Imperial Eagle A. adalberti
Booted Eagle H. pennatus*
Tawny Eagle A. rapax
Little Eagle H. morphnoides
Golden Eagle A. chrysaetos*
African Black Eagle A. verreauxii
Wedge-tailed Eagle A. audax
Gurney's Eagle A. gurneyi
Bonelli's Eagle A.fasciata
African Hawk-Eagle A. spilogaster
Cassin's Hawk-Eagle A. africana
Pygmy Eagle H. weiskei
With fasciatus and spilogaster removed from Hieraaetus and placed in Aquila, and with
kienerii removed to the monotypic genus Lophotriorchis, as recommended by both Helbig
et al. (2005) and Lerner & Mindell (2005), the characters now hold for all species presently
included in Hieraaetus and Aquila (Table 2).
Roulin & Wink (2004) evaluated raptors taxonomically by whether they are
monomorphic or polymorphic. They listed three members of present-day Hieraaetus as
polymorphic, but Bonelli's Eagle as monomorphic. The latter is now included in Aquila, as
advocated by Lerner & Mindell (2005) and Helbig et al. (2005), and the other two taxa in
the revamped Hieraaetus are also polymorphic, thus the results of Roulin & Wink (2004) are
consistent with retention of Hieraaetus.
Debus et al. (2007b), in discussing the breeding biology of Little Eagle H. morphnoides,
stated The results of this study and that on the Wedge-tailed Eagle (Debus et al. 2007a)
lend some support to the separation of the genera Hieraaetus and Aquila, in the revised
sense (following Helbig et al. 2005 and Lerner & Mindell 2005)/ Further, Debus (2011)
concluded 'Pair 3 provided an even greater contrast with the Wedge-tailed Eagle, in terms
of parental sex-roles, than did the other Little Eagle pairs in 2006-2008 (cf Debus et al. 2007a,
b). Thus behavioural and other differences support the continued recognition of the two
genera ( Aquila and Hieraaetus), although internationally they tend now to be combined (e.g.
Gjershaug et al. 2009).'
Bunce et al. (2005) analysed the mt DNA of an extinct large eagle from New Zealand
and concluded that it was closer to species in Hieraaetus than those in Aquila. They theorised
that this large eagle evolved from a small Hieraaetus in Australia or Asia. Their results
provide yet another indication that Aquila and Hieraaetus are different.
Ayres' Eagle. It seems strange that H. ayresii has been called Ayres' Hawk-Eagle, the
only one of the revamped genus to be so named. The name 'Hawk-Eagle' refers primarily
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
William S. Clark
297
Bull. B.O.C. 2012 132(4)
to slender, long-tailed, feathered-tarsi eagles that hunt inside forests in the manner of
overgrown goshawks, such as the various members of the Old World genus Nisaetus
(formerly Spizaetus, which is now restricted to four Neotropical species (Helbig et al. 2005)
but also for two African raptors, African Hawk-Eagle Aquila spilogaster and Cassin's Hawk-
Eagle A. africanus. These hawk-eagles have fairly short broad wings and long tails suitable
for hunting inside forests, whereas Ayres' Eagle has rather narrower and longer wings, and
a relatively shorter tail. Ayres' is an aerial hunter in the manner of Booted and Little Eagles,
and shares few characters with true hawk-eagles, which hunt primarily inside forests. For
consistency of nomenclature, it should be called simply 'Ayres' Eagle', as was done many
years ago by Roberts (1940).
Conclusion. Herein I present rationale for retaining the genus Hieraaetus. I also dispute
that any of the references cited by Sanger et al. (2005) justify moving this genus into Aquila
and further dispute that the results of Wink & Sauer-Giirth (2000) justified such inclusion.
Both Aquila and Hieraaetus, as presently constituted (Table 2), are monophyletic and
sister taxa (Helbig et al. 2005, Lerner & Mindell 2005, supported by Griffiths et al. 2007,
Haring et al. 2007), with the spotted eagles placed in their own genus Clanga, which is also
monophyletic but forms a clade with the somewhat morphologically different Lophaetus
and Ictinaetus (Wells & Inskipp 2012). Wells & Inskipp (2012) wrote 'A further advantage
of this smaller genus approach is that it facilitates retention of the name Hieraaetus.' Note
that all DNA-based studies have retained Hieraaetus as a distinct genus (e.g., Griffiths et al.
2007, Haring et al. 2007).
Acknowledgements
R. Banks, S. Debus, V. Remsen, G. Sangster and D. Wells provided helpful comments on earlier drafts, and
R. Dowsett and A. Kemp provided additional comments on the submitted manuscript.
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Griffiths, C. S., Barrowclough, G. F., Groth, J. G. & Mertz, L. A. 2007. Phylogeny, diversity, and classification
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of the hawk-eagles of the genus Spizaetus (Aves, Accipitridae): phylogenetic analyses based on
mitochondrial markers. /. Zoo/. Syst. & Evol. Res. 45: 353-365.
Helbig, A. J., Kocum, A., Seibold, I. & Braun, M. J. 2005. A multi-gene phylogeny of aquiline eagles (Aves:
Acciptriformes) reveals extensive paraphyly at the genus level. Mol. Phyl. & Evol. 35: 147-164.
Hockey, P. A. R., Dean, W. R. J. & Ryan, P. G. (eds.) 2005. Roberts' birds of Southern Africa. Seventh edn.
Trustees of the John Voelker Bird Book Fund, Cape Town.
Lerner, H. R. L. & Mindell, D. P. 2005. Phylogeny of eagles. Old World vultures, and other Accipitridae based
on nuclear and mitochondrial DNA. Mol. Phyl. & Evol. 37: 327-346.
Roberts, A. 1940. The birds of South Africa. H. F. & G. Witherby, London.
Roulin, A. & Wink, M. 2004. Predator-prey relationships and the evolution of colour polymorphism: a
comparative analysis in diurnal raptors. Biol. J. Finn. Soc. 81: 565-578.
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Sangster, G., Collison, J. ML, Helbig, A. J., Knox, A. G. & Parkin, D. T. 2005. Taxonomic recommendations for
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Cl. 132: 70-72.
Address: 2310 South Whitehouse Circle, Harlingen, Texas 78550, USA, e-mail: raptours@earthlink.net
© 2012 The Authors; Journal compilation © 2012 British Ornithologists' Club
Bulletin of the
British Ornithologists’ Club
Edited by Guy M. Kirwan
Index for Volume 132 (2012)
Author and Contents Index
300
Bull. B.O.C. 2012 132(4)
LIST OF AUTHORS AND CONTENTS
ABREU, M. See JIMENEZ, S.
ABENTE, J. S. See AZPIROZ, A. B.
ABENTE, J. S. See JIMENEZ, S.
AGUIAR, K. M. O. See INGELS, J.
AMAKOBE, B. See PEARSON, D.
AMIGO, X. See GREENEY, H. F.
ANGULO PRATOLONGO, F., FLANAGAN, J. N. M., VELLINGA, W.-P. & DURAND, N. Notes on
the birds of Laquipampa Wildlife Refuge, Lambayeque, Peru 162
AZPIROZ, A. B. See JIMENEZ, S.
AZPIROZ, A. B., MENENDEZ, J. L., JARAMILLO, A., PRESA, D„ CALIMARES, C., SARALEGUI, A.
& ABENTE, J. S. New information on the distribution and status of birds in Uruguay 46
BAIKER, J. See FJELDSA, J.
BEEHLER, B. M., DIAMOND, J. M., KEMP, N., SCHOLES, E., MILENSKY, C. & LAMAN, T. G.
Avifauna of the Foja Mountains of western New Guinea 84
BENSCH, S. See PEARSON, D.
BESSON, L. The collecting history and distribution of Dusky Friarbird Philemon fuscicapillus
(Morotai, Indonesia) elucidated from museum data 236
BLACK, A. B. Collection localities of the Night Parrot Pezoporus ( Geopsittacus ) occidentalis
(Gould, 1861) 277
BOSTOCK, N. See KIRWAN, G. M.
BUDEN, D. W. Seabirds of Sorol Atoll, Yap, Federated States of Micronesia 116
BUGONI, L. See KLEIN, S. R.
CALIMARES, C. See AZPIROZ, A. B.
CAMBREZY, C. See CLAESSENS, O.
CARLOS, C. J., ROSELAAR, C. S. & VOISIN, J.-F. A replacement name for Charadrius leschenaultii
crassirostris (Severtzov, 1873), a subspecies of Greater Sand Plover 63
CLAESSENS, O., CAMBREZY, C., COBIGO, M., MAILLE, S. & RENAUDIER, A. So far from
the Andes: Black-and-white Tanager Conothraupis speculigera, an unexpected vagrant to
French Guiana 55
COBIGO, M. See CLAESSENS, O.
CLARK, W. S. The eagle genus Hieraaetus is distinct from Aquila, with comments on the name
Ayres' Eagle 295
DAUDT, N. W. See KLEIN, S. R.
DAVISON, G. W. H., JIANG, C„ ZHENGWANG, Z. & DE, C. Full tree resolution of Polyplectron
Temminck, 1813, confirms species status of Hainan P. katsumatae Rothschild, 1906, and Bornean
Peacock-Pheasants P. schleiermacheri Briiggemann, 1877 251
DE, C. See DAVISON, G. W. H.
DEAN, W. R. J., FRANKE, U., JOSEPH, G„ GONgALVES, F. M., MILLS, M. S. L„ MILTON, S. J.,
MONADJEM, A. & OSCHADLEUS, H. D. Type specimens in the bird collection at
Lubango, Angola 41
DIAMOND, J. M. See BEEHLER, B. M.
DICKINSON, E. C. See GREGORY, S. M. S.
DICKINSON, E. C. See SVENSSON, L.
DICKINSON, E.C. & Svensson, L. A new name for a buzzard from the Himalayas 221
DONATELLI, R. J. See UBAID, F. K.
DONEGAN, T. M. Geographical variation in Immaculate Antbird Myrmeciza immaculata , with a
new subspecies from the Central Andes of Colombia 3
DONEGAN, T. M. Range extensions and other notes on the birds and conservation of the Serrania de
San Lucas, an isolated mountain range in northern Colombia 140
DOWSETT, R. J. Two pre-occupied names in Afrotropical ornithology 69
Author and Contents Index
301
Bull. B.O.C. 2012 132(4)
DURAND, N. See ANGULO PRATOLONGO, F.
ENGBLOM, G. See FJELDSA, J.
FLANAGAN, J. N. M. See ANGULO PRATOLONGO, F.
FJELDSA, J., BAIKER, J., ENGBLOM, G., FRANKE, I., GEALE, D., KRABBE, N. K., LANE,
D. F„ LEZAMA, M„ SCHMITT, F., WILLIAMS, R. S. R„ UGARTE-NUNEZ, J., YABAR, V. &
YABAR, R. Reappraisal of Koepcke's Screech Owl Megascops koepckeae and description of a
new subspecies 180
FRAHNERT, S. See PEARSON, D.
FRAHNERT, S. See SVENSSON, L.
FRANKE, I. See FJELDSA, J.
FRANKE, U. See DEAN, W. R. J.
GARCIA, T. See GREENEY, H. F.
GEALE, D. See FJELDSA, J.
GELIS, R. A. See GREENEY, H. F.
GONgALVES, F. M. See DEAN, W. R. J.
GREENEY, H. F. & JIPA, M. The nest of Crescent-faced Antpitta Grallaricula lineifrons in
north-east Ecuador 217
GREENEY, H. F., GELIS, R. A., GARCIA, T. & AMIGO, X. The nest, eggs and nestlings
of Fulvous Antshrike Frederickena fulva from north-east Ecuador 65
GREGORY, S. M. S. & DICKINSON, E. C. Clanga has priority over Aquiloides (or how to drop
a clanger) 135
GRIEVE, A. & KIRWAN, G. M. Studies of Socotran birds VII. Forbes- Watson's Swift Apus berliozi in
Arabia— the answer to the mystery of the 'Dhofar swift' 194
HORNBUCKLE, J. See KIRWAN, G. M.
HORST, L. See LECROY, M.
INGELS, J., RENAUDIER, A. & AGUIAR, K. M. O. Status of White-bellied Spinetail Synallaxis
propinqua in French Guiana and Amapa (Brazil) 60
INSKIPP, T. See WELLS, D. R.
JANSEN, J. F. J. Missiemuseum, Steijl, the Netherlands— the history of a little-known collection 212
JARAMILLO, A. See AZPIROZ, A. B.
JIANG, C. See DAVISON, G. W. H.
JIMENEZ, S„ ABENTE, J. S„ AZPIROZ, A. B„ SAVIGNY, C. & ABREU, M. First Uruguayan records
of Great-winged Petrel Pterodroma macroptera 209
JIPA, M. See GREENEY, H. F.
JOSEPH, G. See DEAN, W. R. J.
KERTON, I. See PEARSON, D.
KEMP, N. See BEEHLER, B. M.
KIRCHMAN, J. J. See LECROY, M.
KIRWAN, G. M. See GRIEVE, A.
KIRWAN, G. M. See SCHODDE, R.
KIRWAN, G. M., BOSTOCK, N., HORNBUCKLE, J., MARSHALL, A. & OXLADE, M. Does
Gull-billed Tern Gelochelidon nilotica breed in the interior of continental South America? 133
KLEIN, S. R., DAUDT, N. W. & BUGONI, L. Bulwer's Petrel Bulweria bulwerii in Brazilian waters 214
KRABBE, N. K. See FJELDSA, J.
LAMAN, T. G. See BEEHLER, B. M.
LANE, D. F. See FJELDSA, J.
LECROY, M., KIRCHMAN, J. J. & HORST, L. Rediscovery of missing specimens once held at
Vassar College, including the holotype of Baudo Guan Penelope ortoni 222
LEZAMA, M. See FJELDSA, J.
MAFFEI, F. See UBAID, F. K.
MAILLE, S. See CLAESSENS, O.
Author and Contents Index
302
Bull. B.O.C. 2012 132(4)
MANEGOLD, A. On the name of the Canary Blue Tit Cyanistes teneriffae from Gran Canaria 68
MARSHALL, A. See KIRWAN, G. M.
MENENDEZ, J. L. See AZPIROZ, A. B.
MILENSKY, C. See BEEHLER, B. M.
MILLS, M. S. L. See DEAN, W. R. J.
MILTON, S. J. See DEAN, W. R. J.
MONADJEM, A. See DEAN, W. R. J.
MOYA, G. M. See UBAID, F. K.
OSCHADLEUS, H. D. See DEAN, W. R. J.
OXLADE, M. See KIRWAN, G. M.
PATTEN, M. A. The White Ibis Eudocimus albus subspecies of South America 128
PEARSON, D., KERTON, I., AMAKOBE, B. & BENSCH, S. Presumed hybrid Blyth's Reed Acrocephalus
dumetorum x Marsh Warbler A. palustris trapped at Ngulia, Kenya, in December 2009 124
PEARSON, D., SVENSSON, L. & FRAHNERT, S. Further on the type series and nomenclature of the
Isabelline Shrike Lanius isabellinus 270
PORTER, R. See SCHODDE, R.
PRESA, D. See AZPIROZ, A. B.
RENAUDIER, A. See CLAESSENS, O.
RENAUDIER, A. See INGELS, J.
RHEINDT, F. E. New avian records from the little-explored Fakfak Mountains and the Onin Peninsula
(West Papua) 102
ROSELAAR, C. S. See CARLOS, C. J.
SARALEGUI, A. See AZPIROZ, A. B.
SAVIGNY, C. See JIMENEZ, S.
SCHMITT, F. See FJELDSA, J.
SCHODDE, R., KIRWAN, G. M. & PORTER, R. Morphological differentiation and speciation among
darters ( Anhinga ) 283
SCHOLES, E. See BEEHLER, B. M.
SEVILLANO RIOS, C. S. Rainbow Starfrontlet Coeligena iris in Huascaran National Park, Ancash,
Peru 207
SHIRIHAI, H. See SVENSSON, L.
SHIRIHAI, H. Correcting the identification of two rare wheatear records in Israel 226
SVENSSON, L. See PEARSON, D.
SVENSSON, L. A new subspecies of Western Orphean Warbler Sylvia hortensis and criteria for
separating Western from Eastern Orphean Warbler S. crassirostris 75
SVENSSON, L., SHIRIHAI, H., FRAHNERT, S. & DICKINSON, E. C. Taxonomy and nomenclature
of the Stonechat complex Saxicola torquatus sensu lato in the Caspian region 260
UBAID, F. K., MAFFEI, F., MOYA, G. M. & DONATELLI, R. J. Range extension for Buff-fronted Owl
Aegolius harrisii in south-east Brazil 175
UGARTE-NUNEZ, J. See FJELDSA, J.
VELLINGA, W.-P. See ANGULO PRATOLONGO, F.
VOISIN, J.-F. See CARLOS, C. J.
WELLS, D. R. & INSKIPP, T. A proposed new genus of booted eagles (tribe Aquilini) 70
WILLIAMS, R. S. R. See FJELDSA, J.
YABAR, R. See FJELDSA, J.
YABAR, V. See FJELDSA, J.
ZHENGWANG, Z. See DAVISON, G. W. H.
Scientific Names Index
303
Bull. B.O.C. 2011 132(4)
INDEX TO SCIENTIFIC NAMES
All generic and specific names (of birds only) are indexed. New specific and subspecific names are indexed
in bold print under generic, specific and subspecific names. Illustrations and figures are numbered in italics.
abellei, Arremon 174
Aburria aburri 144, 153
aburri, Aburria 144, 153
acadicus, Aegolius 178
Accipiter bicolor 170
Accipiter buergersi 93, 96
Accipiter cirrhocephalus 96
Accipiter meyerianus 93, 96
Accipiter novaehollandiae 96
Accipiter poliocephalus 96, 105
Accpiter striatus 170
Achaetops pycnopygius 43
Acrocephalus baeticatus 124
Acrocephalus dumetorum 124-127, 126
Acrocephalus palustris 124
Acrocephalus scirpaceus 124
Actitis hypoleucos 96
Actitis macularius 170
acutipennis, Chordeiles 171
adalberti, Aquila 72, 296
Adelomyia melanogenys 171
adolphinae, Myzomela 92, 93
aedon. Troglodytes 173
Aegolius acadicus 178
Aegolius funereus 178
Aegolius harrisii 46, 48, 49, 175-179, 176
Aegolius ridgwayi 178
Aegotheles albertisi 91, 97
Aegotheles archboldi 91
Aegotheles insignis 91, 97
Aepypodius arfakianus 96
aequinoctialis, Geothlypis 174
Aeronautes monti vagus 171
Aethopyga saturata 258
affinis, Apus 195, 205
africana, Aquila 71, 72, 296
africana, Cassinaetus 72
africana, Hieraaetus 72
africana, Mirafra 42
africana, Spizaetus 72
africanus, Aquila 297
africanus, Spizaetus 70
Aglaeactis cupripennis 171
Ailuroedus buccoides 94, 101, 107
Ailuroedus melanotis 94, 101, 107
alba, Ardea 170
alba, Gygis 121
alba, Scolopax 129
alba, Tyto 171, 177
albertisi, Aegotheles 91, 97
albertisi, Drepanornis 92, 101
albertisi, Gymnophaps 88
albertisii, Gymnophaps 97
albescens, Synallaxis 157
albiceps, Atlapetes 174
albicollis, Nyctidromus 171
albifrons, Henicophaps 105
albinucha, Atlapetes 157
albipennis, Penelope 162, 164
albispecularis, Poecilodryas 88, 92, 99
albitarsis, Ciccaba 171
alboauricularis, Lichmera 102, 112, 114
albogriseus, Pachyramphus 173
albogularis. Megascops 182, 183, 186, 188, 193
albolimbata, Rhipidura 91, 94, 99
albonotata, Meliphaga 102, 112, 114
alboscapulatus, Malurus 98
albus, Eudocimus 128-132, 130-131
albus ramobustorum subsp. nov., Eudocimus 128
Alcedo azurea 98
Alcedo pusilla 105
alecto, Myiagra 99, 106
Alectoris chukar 254
Alisterus amboinensis 105
Alisterus chloropterus 97
alixii, Clytoctantes 140, 153
Amalocichla incerta 88, 92, 99
Amaurospiza concolor 165, 167, 174
amazilia, Amazilia 171
Amazilia amazilia 171
Amazilia castaneiventris 140, 153
Amazona farinosa 154
Amblyornis flavifrons 85, 87, 89-90, 92, 94, 101
Amblyornis inornatus 92, 102, 113-114
Amblyornis macgregoriae 92
amboinensis, Alisterus 105
amboinensis, Macropygia 96, 105
americana, Chloroceryle 171
Amytornis goyderi 280
Anabacerthia 152
anais, Mino 101, 107
analoga, Meliphaga 94, 100, 107
andrei, Chaetura 156
angolensis, Mirafra 43
Anhinga 283-294
anhinga, Anhinga 283, 286, 289-292
Anhinga anhinga 283, 286, 289-292
Anhinga melanogaster 283-294
Anhinga [melanogaster] novaehollandiae 293
Anhinga novaehollandiae 283-294
Anhinga rufa 283-294
Anous minutus 120
Anous stolidus 120
anthonyi, Caprimulgus 171
Anthus bogotensis 173
antisiensis, Cranioleuca 172
Aplonis metallica 101, 107
Aplonis opaca 122
Apus affinis 195, 205
apus, Apus 194-199, 200, 205
Apus apus 194-199, 200, 205
Apus balstoni 194
Apus barbatus 194
Apus berliozi 194-206, 201, 202, 204, 205
Apus bradfieldi 194
Apus horns 42
Apus melba 195
Apus niansae 195
Apus pallidus 194-199, 200
Apus toulsoni 42
Scientific Names Index
304
Bull. B.O.C. 2012 132(4)
Apus unicolor 194
Aquila 70-72, 295-298
Aquila adalberti 72, 296
Aquila africana 71-72, 296
Aquila africanus 297
Aquila audax 72, 296
Aquila chrysaetos 296
Aquila clanga 70-71, 135-136, 213, 295
aquila, Eutoxeres 140, 143, 144
Aquila fasciata 71-72, 296
Aquila gumeyi 72, 296
Aquila hastata 70-71, 135, 295
Aquila heliaca 296
Aquila nipalensis 72, 296
Aquila pomarina 70-71, 135, 295
Aquila rapax 72, 296
Aquila spilogaster 71-72, 296, 297
Aquila verreauxii 72, 296
Aquila wahlbergi 295
Aquiloides 135-136
Aquiloides, gen. nov. 71
Aramides axillaris 165, 170
Aratinga erythrogenys 170
Aratinga wagleri 170
archboldi, Aegotheles 91
arctica, Fratercula 213
Ardea alba 170
Ardea sumatrana 108
arfakianus, Aepypodius 96
arfakianus, Sericornis 89, 91, 93, 98
arfaki, Oreocharis 92, 100
ariel, Fregata 119
arminjoniana, Pterodroma 211
Arremon abellei 174
Arremon assimilis 174
Arses insularis 99
Arses telescophthalmus 106
Artamus maximus 101
arthus, Tangara 140, 148
aruensis, Meliphaga 100
aspasia, Leptocoma 107
aspasia, Nectarinia 100
assimilis, Arremon 174
Astrapia 93
ater, Manucodia 101
ater, Melipotes 92
aterrimus, Probosciger 97, 105
Athene cunicularia 171
Atlapetes albiceps 174
Atlapetes albinucha 157
Atlapetes leucopterus 174
Atlapetes seebohmi 174
atra, Chalcopsitta 105
atra, Rhipidura 91, 94, 99
atratus, Coragyps 170
atratus, Scytalopus 140, 146, 157
atricaudus, Myiobius 155
atrifrons, Zosterops 94, 100
atripennis, Saltator 15
atrogularis, Prinia 258
atrovirens, Lalage 98, 106
audax, Aquila 72, 296
aura, Cathartes 170
aurantiirostris, Catharus 140, 148, 157
aureliae, Haplophaedia 157
aureus, Sericulus 101
auriceps, Pharomachrus 140, 144
auriculata, Zenaida 170
aurulentus, Piculus 50
australis, Tchagra 44
Aviceda subcristata 96
axillaris, Aramides 165, 170
axillaris, Monarcha 91, 99, 110
ayresii, Hieraaetus 72, 296
azarae, Synallaxis 157, 172
azurea, Alcedo 98
baeticatus, Acrocephalus 124
bairdii, Myiodynastes 173
balstoni, Apus 194
barbata, Penelope 164
barbatus, Apus 194
barbatus, Pycnonotus 78
Basileuterus fraseri 174
Basileuterus nigrocristatus 174
Basileuterus trifasciatus 174
Basileuterus tristriatus 20, 157
Batara 66
Batara cinerea 66
Batis molitor 43
beccarii, Gallicolumba 91, 96
belfordi, Melidectes 93
belicosa, Sturnella 174
bennetti, Casuarius 87, 96, 213
bergii, Thalasseus 116, 121
berigora, Falco 96
berlepschi, Myrmeciza 4, 8, 19, 21, 22
berlepschi, Parotia 87, 90, 92, 94, 101
berliozi, Apus 194-206, 200-202 , 204-205
bernardi, Thamnophilus 172
bernsteini, Centropus 97, 105
bernsteini. Sterna 213
bicalcaratum, Polyplectron 251-254, 255, 257
bicolor, Accipiter 170
bicolor, Tiaris 140, 150
bimaculata, Peneothello 93, 99
blainvillii, Peltops 94, 101, 106
bogotensis, Anthus 173
bolivianum, Glaucidium (jardinii) 190
bombus, Chaetocercus 166, 171
bonapartei, Nothocercus 157
bonariensis, Molothrus 174
bonariensis, Thraupis 173
boyeri, Coracina 98, 106
brachyrhyncha, Rhipidura 91, 99
brachyura, Chaetura 171
brachyura, Poecilodryas 99
bracteatus, Dicrurus 101, 107
bradfieldi, Apus 194
brasilianum, Glaucidium 49, 177
brevicauda, Muscigralla 173
brevipes, Monticola 43
brevirostris, Lugensa 209, 210, 211
bruijnii, Drepanornis 101
bruijnii, Grallina 101
bruijnii, Micropsitta 91, 93, 97
brunnescens, Premnoplex 157
Bubo virginianus 177
Bubulculus ibis 170
buccoides, Ailuroedus 94, 101, 107
buceroides, Philemon 100, 107
Scientific Names Index
305
Bull. B.O.C. 2011 132(4)
buergersi, Accipiter 93, 96
Bulweria 211
Bulweria bulwerii 214-216, 215
Bulweria fallax 215
bulwerii, Bulweria 214-216, 215
Buphagus erythrorhynchus 44
Burhinus superciliaris 170
burmanicus, Buteo 221
Busarellus nigricollis 46, 48
Buteo 221
Buteo burmanicus 221
Buteo (buteo) hodgsoni nom. nov. 221
Buteogallus meridionalis 170
Buteogallus umbitinga 170
Buteo lagopus 221
Buteo leucorrhous 170
Buteo polyosoma 170
Buteo refectus 221
Butorides striata 170
Cacatua galerita 97, 105
Cacomantis 88
Cacomantis castaneiventris 97, 105
Cacomantis variolosus 97, 105
caerulea, Egretta 170
caeruleogrisea, Coracina 98, 106
caerulescens, Ptilorrhoa 93, 98
Caliechthrus leucolophus 97
callonotus, Verniliornis 172
calopterus, Mecocerculus 172
Campephilus gayaquilensis 172
Campochaera sloetii 98
Camptostoma obsoletum 172
Campylorhamphus pusillus 157
Campylorhamphus trochilirostris 152, 157
Campylorynchus fasciatus 173
Cantor chilus superciliaris 173
canus, Picus 258
capensis, Zonotrichia 174
capistratus, Serinus 44
Capito hypoleucus 23
Caprimulgus anthonyi 171
Caprimulgus cayannensis 150
Caprimulgus longirostris 50, 53
Caprimulgus macrurus 97
Caracara cheriway 170
Carduelis magellanica 174
Carduelis psaltria 174
caripensis, Steatornis 166, 171
carolae, Melipotes 92, 100
carolae, Parotia 92
cassicus, Cracticus 101, 107
Cassinaetus africana 72
castaneiventris, Amazilia 140, 153
castaneiventris, Cacomantis 97, 105
castanonota, Ptilorrhoa 93, 98, 106
Casuarius bennetti 87, 96, 213
casuarius, Casuarius 104
Casuarius casuarius 104
Casuarius unappendiculatus 96, 213
Catamblyrhynchus diadema 173
Cathartes aura 170
Catharus aurantiirostris 140, 148, 157
Catharus dryas 157
Catharus fuscater 173
Catoptrophorus semipalmatus 46, 48
caudata, Inezia 150
cayannensis, Caprimulgus 150
centralis. Megascops 188
centralis, Megascops 'guatemalae' 140, 149
centralis, Megascops (guatemalae) 182
centralis. Megascops [guatemalae] 187, 193
Centropus bernsteini 97, 105
Centropus menbeki 97, 105
Ceyx lepidus 98, 105
Chaetocercus bombus 166, 171
Chaetorhynchus papuensis 101, 107
Chaetura andrei 156
Chaetura brachyura 171
Chaetura chapmani 156
chalconota, Ducula 88, 91
Chalcophaps stephani 96, 105
Chalcopsitta atra 105
Chalcopsitta duivenbodei 97
chalcurum, Polyplectron 251-253, 254, 257
chalybata, Manucodia 108
chalybatus, Manucodia 101
chalybea, Progne 173
chapmani, Chaetura 156
Charadrius 64
Charadrius crassirostris 64
Charadrius leschenaultii 63-65
Charadrius leschenaultii scythicus nom. nov. 64
Charadrius vociferus 170
Charadrius wilsonia 64
Charmosyna josefinae 97
Charmosyna placentis 93, 97, 105
Charmosyna pulchella 93, 97
cheriway, Caracara 170
chiguanco, Turdus 173
chiniana, Cisticola 43
Chloroceryle americana 171
Chloroceryle inda 140, 150
chloronota, Gerygone 89, 99
chloropterus, Alisterus 97
Chlorospingus flavigularis 140, 148
choliba. Megascops 177, 182, 184, 185, 187, 188, 193
choliba, Otus 183, 185
Chordeiles acutipennis 171
chrysaetos, Aquila 296
Chrysococcyx meyeri 93, 97
Chrysococcyx minutillus 93, 97
Chrysococcyx ruficollis 88, 91, 93, 97
chrysogaster, Gerygone 99, 110
chrysogaster, Pheucticus 174
chrysomela, Monarcha 99, 106
chrysoptera, Daphoenositta 102, 111, 114
chukar, Alectoris 254
Ciccaba albitarsis 171
Cicinnurus magnificus 94, 101, 107
Cicinnurus regius 101, 107
cincta, Riparia 43
cinerea, Batara 66
cinerea, Gerygone 89, 101
cinerea, Motacilla 98
cinerea, Piezorhina 174
cinereum, Conirostrum 173
cinereum, Todirostrum 172
cinereus, Contopus 172
Cinnyris fuscus 44
Cinnyris jugularis 107
Scientific Names Index
306
Bull. B.O.C. 2012 132(4)
Circus plumipes 221
cirrhocephalus, Accipiter 96
Cisticola chiniana 43
Cisticola subruficapilla 43
citrinelloides, Serinus 44
Clanga 135-136,297
clanga, Aquila 70-71, 135, 136, 213, 295
clanga, Clanga 136
Clanga clanga 136
Clanga fasciata 135
Clanga macrodactyla 135
Clanga naevia 135
Clytoceyx rex 98
Clytoctantes alixii 140, 153
Coccyzus erythropthalmus 171
coco, Tantalus 129
coelestis, Forpus 171
Coeligena iris 171, 207, 207-209, 208
Coereba flaveloa 174
coerulescens, Saltator 156
Colaptes rivolii 140, 144
Colaptes rubiginosus 172
Colibri coruscans 171
Collocalia esculenta 98
Collocalia hirundinacea 98
Collocalia nitens 106
Collocalia vanikorensis 98, 106
Colluricincla megarhyncha 100, 107
collurio, Lanius 270-271, 274
colombica, Thalurania 140, 150, 152
Columba livia 170
columbiana, Ortalis 156, 157
Columbina cruziana 170
concepcion subsp. nov., Myrmeciza immaculata
11
concolor, Amaurospiza 165, 167, 174
concolor, Corythaixoides 42
Conirostrum cinereum 173
Conothraupis mesoleuca 55
Conothraupis speculigera 55-59, 56, 173
conspicillata, Procellaria 211
contaminatus, Fleliobletus 46, 48, 50
Contopus cinereus 172
Contopus fumigatus 172
Coracina 112
Coracina boyeri 98, 106
Coracina caeruleogrisea 98, 106
Coracina incerta 98
Coracina lineata 88, 101
Coracina melaena 98
Coracina melas 106
Coracina montana 91, 93, 98
Coracina morio 106
Coracina papuensis 98, 106
Coracina schisticeps 93, 98
Coragyps atratus 170
coronulatus, Ptilinopus 102, 109, 114
coruscans, Colibri 171
Corvus fuscicapillus 112
Corvus tristis 101, 107
Corythaixoides concolor 42
Cracticus cassicus 101, 107
Cracticus quoyi 101, 107
Cranioleuca antisiensis 172
Cranioleuca erythrops 140, 144, 146
crassirostris, Charadrius 64
crassirostris, Eudromias 63, 64
crassirostris, Rhamphocharis 90, 92, 100
crassirostris, Sylvia 75-83, 77, 80
Crateroscelis murina 93, 98, 106
Crateroscelis nigrorufa 88, 101
Crateroscelis robusta 88, 91, 93, 98
cristata, Goura 108
cristatus, Oxyruncus 152, 157
cristatus, Pitohui 100, 107
Crithagra gularis 69
Crithagra scotops 69
Crithagra scotops kirbyi, nom. nov. 69
Crotophaga sulcirostris 171
cruentata, Myzomela 93, 100, 107
cruziana, Columbina 170
cryptoleuca, Peneothello 89, 92, 93, 99
cunicularia, Athene 171
cupripennis, Aglaeactis 171
cuvieri, Talegalla 108
Cyanistes teneriffae 68
cyanocephala, Euphonia 48, 53
cyanocephalus, Malurus 98, 106
Cyanocorax mystacalis 173
Cyanocorax yncas 157
cyanoleuca, Pygochelidon 173
Cyanopsitta spixii 213
cyanus, Peneothello 92, 93, 99
Cyclarhis gujanensis 173
Cyclopsitta diophthalma 109
Cyclopsitta gulielmitertii 97, 102, 109, 114
Cymbilaimus 66-67
Cymbilaimus lineatus 66, 155
Cymbilaimus sanctaemariae 66
cyrenaicae, subsp. nov., Sylvia hortensis 79
Dacelo gaudichaud 98, 106
dactylatra, Sula 116, 120
Daphoenositta chrysoptera 102, 111, 114
dauma, Zoothera 98
decollatus, Megapodius 96
Dendrocolaptes platyrostris 46, 48, 50
Dendrocopos medius 213
Dendropicos fuscescens 69
Dendropicos namaquus 69
desmarestii, Psittaculirostris 105
diadema, Catamblyrhynchus 173
Dicaeum pectorale 100, 107
dichrous, Pitohui 94, 100, 111
Dicrurus bracteatus 101, 107
Diglossa sittoides 173
diophthalma, Cyclopsitta 109
Dives warszewiczi 174
Drepanornis albertisi 92, 101
Drepanomis bruijnii 101
Dromococcyx pavoninus 140, 143
dryas, Catharus 157
Drymodes supercilaris 99
Dryocopus lineatus 172
Ducula 84, 96
Ducula cf. chalconota / sp. nov. 88
Ducula chalconota 88, 91
Ducula pinon 97, 105
Ducula rufigaster 97, 105
Ducula zoeae 97, 105
duivenbodei, Chalcopsitta 97
307
Bull. B.O.C. 2011 132(4)
Scientific Names Index
dumetorum, Acrocephalus 124-127, 126
dumontii, Mino 101, 107
Eclectus roratus 97, 105
Egretta caerulea 170
Egretta rufescens 128
Egretta thula 170
Elaenia flavogaster 52
Elaenia mesoleuca 51
Elaenia spectabilis 51
elegans, Melanopareia 172
ellisiana, Pyriglena 7
Emberizoides herbicola 52
Emberizoides ypiranganus 52
Embernagra platensis 52
Epimachus fastosus 87, 92, 94, 101
Epimachus meyeri 92
episcopus, Thraupis 173
erythrocephala, Pipra 155
erythrocephalus, Hylocryptus 166, 172
erythrogaster. Pitta 98, 106
erythrogenys, Aratinga 170
erythrops, Cranioleuca 140, 144, 146
erythrops, Odontophorus 144, 154
erythropthalmus, Coccyzus 171
erythropygius, Xiphorhynchus 140, 144, 146, 157
erythrorhynchus, Buphagus 44
Erythrura trichroa 100
esculenta, Collocalia 98
Eudocimus 128
Eudocimus albus 128-132, 130
Eudocimus albus ramobustorum subsp. nov. 128
Eudocimus ruber 128
Eudromias crassirostris 63, 64
Eudynamys scolopaceus 97
Eugerygone rubra 92, 99
Euphonia cyanocephala 48, 53
Euphonia laniirostris 174
Eurostopodus papuensis 98
Eurystomus orientalis 106
Euscarthmus meloryphus 172
Eutoxeres aquila 140, 143, 144
exilis, Ixobrychus 170
exsul, Myrmeciza 7-8, 21
fagani. Prunella 204
Falco berigora 96
Falco maculatus 135-136
Falco peregrinus 170
Falco plumipes 221
Falco sparverius 170
fallax, Bulweria 215
fallax, Glycichaera 107
fannyi, Thalurania 15, 140, 150, 152
fanny, Myrtis 171
farinosa, Amazona 154
fasciata, Aquila 71-72, 296
fasciata, Clanga 135
fasciata, Hieraaetus 72
fasciata, Patagioenas 170
fasciatus, Campylorynchus 173
fasciatus, Myiophobus 172
fastosus, Epimachus 87, 92, 94, 101
ferrugineipectus, Grallaricula 167, 172, 219
ferrugineus, Pitohui 100, 107
finschii, Oenanthe 226, 228, 230-231, 234
flava, Piranga 174
flaveloa, Coereba 174
flaveola, Sicalis 174
flavifrons, Amblyornis 85, 87, 89-90, 92, 94, 101
flavigularis, Chlorospingus 140, 148
flavirictus, Meliphaga 102, 112, 114
flavirostris, Grallaricula 218
flavi venter, Machaerirhynchus 94, 99, 106
flaviventer, Xanthotis 100, 107, 112
flavogaster, Elaenia 52
flavogriseum, Pachycare 92, 94, 99
flavovirescens, Microeca 99, 107
forbesi, Rallicula 91
Forpus coelestis 171
fortis, Myrmeciza 4-5, 10, 13, 14, 16, 17, 18, 26,
30-35
Francolinus squamatus 41
fraseri, Basileuterus 174
Fratercula arctica 213
f rater, Monarcha 99, 106
Frederickena 66
Frederickena fulva 65-68
Frederickena unduligera 65
Fregata ariel 119
Fregata minor 118
frontalis, Hemispingus 157
fulgidus, Psittrichas 97
fuliginosus, Tiaris 140, 150, 151
fulva, Frederickena 65-68
fulvigula, Timeliopsis 90, 92, 100
fumigatus, Contopus 172
fumigatus, Melipotes 92, 113
fumigatus, Picoides 140, 144, 172
funereus, Aegolius 178
Furnarius leucopus 172
fusca, Nectarinia 44
fusca, Scolopax 129
fuscata, Onychoprion 121
fuscata, Pseudeos 88, 97
fuscater, Catharus 173
fuscater, Turdus 157, 173
fuscescens, Dendropicos 69
fuscicapilla, Philemon 246
fuscicapilla, Zosterops 100
fuscicapillus, Corvus 112
fuscicapillus, Microphilemon 248
fuscicapillus, Philemon 236-250, 244-245
fuscicapillus, Tropidorhynchus 248
fuscicapillus, Zosterops 92, 94, 111
fuscipenne, Philydor 154
fuscirostris, Talegalla 102, 104, 108, 114
fuscus, Cinnyris 44
galatea, Tanysiptera 98, 105
galerita, Cacatua 97, 105
Gallicolumba beccarii 91, 96
Gallicolumba rufigula 96
Gallinago 96
Galloperdix lunulata 252, 254
Gampsonyx swainsonii 170
garrula, Ortalis 154, 157
gaudichaud, Dacelo 98, 106
gayaquilensis, Campephilus 172
Gelochelidon nilotica 233, 133-135
geoffroyi, Geoff royus 93, 97, 105
Geoff royus geoffroyi 93, 97, 105
Geoffroyus simplex 93, 97
Scientific Names Index
308
Bull. B.O.C. 2012 132(4)
Geothlypis aequinoctialis 174
Geotrygon montana 165, 166
Geranoaetus melanoleucus 170
germaini, Polyplectron 252-253, 254-255, 257
Gerygone chloronota 89, 99
Gerygone chrysogaster 99, 110
Gerygone cinerea 89, 101
Gerygone magnirostris 99, 106
Gerygone palpebrosa 99, 106
Gerygone ruficollis 99, 110
gigantea, Melampitta 113
gigas, Patagona 171
Glaucidium brasilianum 49, 177
Glaucidium (brasilianum) ridgwayi 149
Glaucidium griseiceps 140, 149
Glaucidium hardyi 149
Glaucidium (jardinii) bolivianum 190
Glaucidium peruanum 171, 190
Glycichaera fallax 107
goeldii, Myrmeciza 3-6, 10, 13, 14, 16, 17, 18, 19,
26, 31-35
Goura cristata 108
Goura victoria 96, 213
goyderi, Amytornis 280
graceannae. Icterus 174, 223
Grallaria guatimalensis 172
Grallaria hypoleuca 157
Grallaria ruficapilla 172
Grallaria watkinsi 172
Grallaricula ferrugineipectus 167, 172, 219
Grallaricula flavirostris 218
Grallaricula lineifrons 217-218
Grallaricula nana 219
Grallaricula peruviana 218
Grallina bruijnii 101
gravis, Puffinus 209
grayi, Malurus 98
grimwoodi, Macronyx 43
griseiceps, Glaucidium 140, 149
griseiceps, Pachycephala 107
griseigula, Timeliopsis 102, 112, 114
griseipectus, Lathotriccus 167, 172
griseisticta, Muscicapa 99
griseogularis, Phaethomis 171
griseus, Nyctibius 171
gryphus, Vultur 170
guatimalensis, Grallaria 172
guisei, Ptiloprora 92
gujanensis, Cyclarhis 173
gujanensis, Synallaxis 60
gularis, Crithagra 69
gulielmitertii, Cyclopsitta 97, 102, 109, 114
gurneyi, Aquila 72, 296
guttatus, Hypoedaleus 66
guttula, Monarcha 106
guttulus, Monarcha 99
gutturalis, Habia 23, 156
Gygis alba 121
Gymnocichla 10, 11, 21
Gymnophaps albertisi 88
Gymnophaps albertisii 97
gymnops, Melipotes 92, 113
Habia gutturalis 23, 156
haematodus, Trichoglossus 97, 105
Haematortyx 254
Haematortyx sanguiniceps 252, 254
Halcyon megarhyncha 88, 91, 101
Halcyon nigrocyanea 98
Halcyon sancta 105
Halcyon torotoro 98, 109
Haliastur indus 96, 105
Haplophaedia aureliae 157
hardyi, Glaucidium 149
Harpyhaliaetus solitarius 170
Harpy opsis novaeguineae 96, 108
harrisii, Aegolius 46, 48, 49, 175-179, 176
hastata, Aquila 70-71, 135, 295
hattamensis, Pachycephalopsis 99
heliaca, Aquila 296
Heliobletus contaminatus 46, 48, 50
Heliomaster longirostris 171
Hemiprocne mystacea 98, 106
Hemispingus frontalis 157
Hemispingus melanotis 173
hemprichii, Saxicola 267, 268
Henicopernis longicauda 96
Henicophaps albifrons 105
Henicorhina leucophrys 157-158
Henicorhina leucosticta 156-157
herbicola, Emberizoides 52
Herpsilochmus rufimarginatus 155
Hieraaetus 70, 72, 295-298
Hieraaetus africana 72
Hieraaetus ayresii 72, 296
Hieraaetus fasciata 72
Hieraaetus morphnoides 72, 96, 295-296
Hieraaetus pennatus 70, 72, 295-296
Hieraaetus wahlbergi 72, 295-296
Hieraaetus weiskei 72, 296
hirundinacea, Collocalia 98
Hirundo rustica 173
Hirundo tahitica 98
hispanica, Oenanthe 230
hispaniolensis, Poospiza 174
hockingi, subsp. nov.. Megascops koepckeae 189
hodgsoni nom. nov., Buteo (buteo) 221
hortensis cyrenaicae, subsp. nov., Sylvia 79
hortensis, Sylvia 75-83, 77, 80
horns, Apus 42
hoyi. Megascops 182, 184, 187, 193
Hydropsalis (Caprimulgus) maculicaudus 150
Hydropsalis maculicaudus 140
Hyloclistes virgatus 140, 144, 146
Hylocryptus erythrocephalus 166, 172
hyperythra, Myrmeciza 10
hyperythra, Pachycephala 94, 100, 107, 111
hyperythra, Rhipidura 94, 99, 110
Hypocnemis peruviana 4
Hypoedaleus 66
Hypoedaleus guttatus 66
hypoleuca, Grallaria 157
hypoleuca, Peocilodryas 99, 107
hypoleucos, Actitis 96
hypoleucus, Capito 23
hypopyrogaster, Hypopyrrhus 157
Hypopyrrhus hypopyrogaster 157
ibis, Bubulculus 170
Ibis longirostris 129
Icterus graceannae 174, 223
Icterus mesomelas 174
Scientific Names Index
309
Bull. B.O.C. 2011 132(4)
Ictinaetus 70-71, 295, 297
Ictinaetus malayensis 70
ignobilis, Turdus 140, 148
iliolophum, Oedistoma 100
immaculata concepcion subsp. nov., Myrmeciza
11
immaculata, Myrmeciza 3^0, 13-14, 16, 18-20
immaculatus, Thamnophilus 7
incerta, Amalocichla 88, 92, 99
incerta, Coracina 98
incerta, Pterodroma 211
inda, Chloroceryle 140, 150
indus, Haliastur 96, 105
Inezia caudata 150
inopinatum, Polyplectron 251-253, 254, 257-258
inornatus, Amblyornis 92, 102, 113-114
insignis, Aegotheles 91, 97
insularis. Arses 99
iozonus, Ptilinopus 96, 105
Iridosomis porphyrocephala 157
iris, Coeligena 171, 207, 207-209, 208
isabellinus, Lanius 270-276, 272
isidorei, Pomatostomus 98, 107, 112
Ixobrychus exilis 170
ixoides, Pycnopygius 100
Jabiru mycteria 148
jacarina, Volatina 174
jobiensis, Talegalla 96
josefinae, Charmosyna 97
jugularis, Cinnyris 107
Jynx ruficollis 42
katsumatae, Polyplectron 251-259, 253-254
keiensis, Micropsitta 105
keraudrenii, Manucodia 101, 108
kienerii, Lophotriorchis 296
kirbyi, nom. nov., Crithagra scotops 69
kirhocephalus, Pitohui 94, 100, 111
koepckeae hockingi, subsp. nov.. Megascops 189
koepckeae. Megascops 166, 171, 180-193, 181, 183,
185, 186, 187
koepckei. Megascops 183
laemosticta, Myrmeciza 21
lagopus, Buteo 221
Lalage atrovirens 98, 106
laniirostris, Euphonia 174
Lanioturdus torquatus 43
Lanius collurio 270-271, 274
Lanius isabellinus 270-276, 272
Lanius ruficaudus 272
Lathotriccus griseipectus 167, 172
lawesii, Parotia 92
leachii, Mackenziaena 46, 48, 50
Lepidocolaptes souleyetti 172
lepidus, Ceyx 98, 105
Leptocoma aspasia 107
Leptotila ochraceiventris 166, 170
Leptotila verreauxi 170
Lesbia nuna 171
leschenaultii, Charadrius 63-65
Leucippus sp. 171
leucogaster, Sula 120
leucolophus, Caliechthrus 97
leuconota, Pyriglena 14
leucophrys, Henicorhina 157-158
leucophrys, Vireo 140, 148
leucops, Tregellasia 99, 110
leucoptera, Piranga 157
leucopterus, Atlapetes 174
leucopus, Furnarius 172
leucopyga, Oenanthe 226, 232
leucorrhous, Buteo 170
leucospila, Rallicula 91
leucospodia, Pseudelaenia 172
leucosticta, Ptilorrhoa 93
leucosticta, Henicorhina 156-157
leucosticta, Ptilorrhoa 88, 91, 98
leucostigma, Percnostola 8
leucothorax, Rhipidura 99, 106
leucura, Oenanthe 226- 227, 228
leucura, Saxicola 234
Lichenostomus obscurus 100
Lichmera alboauricularis 102, 112, 114
lineata, Coracina 88, 101
lineatus, Cymbilaimus 66, 155
lineatus, Dryocopus 172
lineifrons, Grallaricula 217-218
Lipaugus unirufus 155, 157
Lipaugus weberi 157
livia, Columba 170
Lonchura 93
Lonchura tristissima 101
longicauda, Henicopernis 96
longicauda, Melanocharis 93, 100
longicaudatus, Mimus 173
longipes, Myrmeciza 10
longirostris, Caprimulgus 50, 53
longirostris. Heliomaster 171
longirostris. Ibis 129
Lophaetus 71, 295, 297
Lophaetus occipitalis 70
Lophorina superba 84, 92-93
Lophotriorchis 72
Lophotriorchis kienerii 296
Lorius lory 97, 105
lory, Lorius 97, 105
Lugensa brevirostris 209, 210, 211
lugens, Oenanthe 226, 228, 230-232
lunulata, Galloperdix 252, 254
Luscinia megarhynchos 79
lyra, Uropsalis 140, 143
macconnelli, Synallaxis 60
macgregoriae, Amblyornis 92
Machaerirhynchus flavi venter 94, 99, 106
Machaerirhynchus nigripectus 91, 94, 99
Mackenziaena 66
Mackenziaena leachii 46, 48, 50
Mackenziaena severa 66
macrodactyla, Clanga 135
Macronyx grimwoodi 43
macroptera, Pterodroma 209-212, 210
Macropygia amboinensis 96, 105
Macropygia nigrirostris 96
macrorrhina, Melidora 98, 105
macrurus, Caprimulgus 97
macularius, Actitis 170
maculatus, Falco 135-136
maculatus, Myiodynastes 173
maculicaudus, Hydropsalis 140
maculicaudus, Hydropsalis (Caprimulgus) 150
maculipectus, Rhipidura 106
Scientific Names Index
310
Bull. B.O.C. 2012 132(4)
madaraszi, Psittacella 88, 91, 97
magellanica, Carduelis 174
magnificus, Cicinnurus 94, 101, 107
magnificus, Ptilinopus 96, 105
magnificus, Ptiloris 101, 108
magnirostris, Gerygone 99, 106
major, Taraba 66
major, Tinamus 153
malacense, Polyplectron 251, 253, 254, 255, 256,
257-258
malayensis, Ictinaetus 70
Malurus alboscapulatus 98
Malurus cyanocephalus 98, 106
Malurus grayi 98
manadensis, Monarcha 99, 106
Manucodia ater 101
Manucodia chalybata 108
Manucodia chalybatus 101
Manucodia keraudrenii 101, 108
marginatus, Microcerculus 156
marshalli. Megascops 186
maurus, Saxicola 260, 261, 262, 264, 266, 267, 268
maximus, Artamus 101
mayri, Ptiloprora 92, 100
mayri, Rallicula 91, 96
Mearnsia novaeguineae 98, 102, 109, 114
Mecocerculus calopterus 172
medius, Dendrocopos 213
Megaceryle torquata 171
Megapodius decollatus 96
Megapodius reinwar dt 105
megarhyncha, Colluricincla 100, 107
megarhyncha. Halcyon 88, 91, 101
megarhynchos, Luscinia 79
megarhynchus, Melilestes 100, 107
Megarhynchus pitangua 52
Megascops albogularis 182-183, 186, 188, 193
Megascops centralis 188
Megascops choliba 177, 182, 184-185, 187, 188, 193
Megascops 'guatemalae' centralis 140, 149
Megascops (guatemalae) centralis 182
Megascops [guatemalae] centralis 187, 193
Megascops (guatemalae) napensis 182
Megascops [guatemalae] napensis 187, 193
Megascops hoyi 182, 184, 187, 188, 193
Megascops koepckeae 166, 171, 180-193, 181, 183,
185, 186, 187
Megascops koepckeae hockingi, subsp. nov. 189
Megascops koepckei 183
Megascops marshalli 186
Megascops napensis 188
Megascops roboratus 171, 182-184, 187-188, 193
Megascops sanctaecatarinae 48
melaena, Coracina 98
Melampitta gigantea 113
melancholicus, Tyrannus 173
Melanerpes pulcher 154
melanoceps, Myrmeciza 3-6, 10, 13, 14, 16-18, 19,
22, 26, 31-35
Melanocharis longicauda 93, 100
Melanocharis nigra 93, 100, 107
Melanocharis versteri 90, 92-93, 100
melanogaster, Anhinga 283-294
melanogenys, Adelomyia 171
melanoleucos, Phalacrocorax 105
melanoleucus, Geranoaetus 170
melanoleucus, Seleucidis 101, 107
melanonota, Pipraeidea 173
Melanopareia elegans 172
melanopis, Theristicus 130
melanops, Turdoides 44
melanotis, Ailuroedus 94, 101, 107
melanotis, Hemispingus 173
melanotos, Sarkidiornis 164
melas, Coracina 106
melba, Apus 195
Melidectes belfordi 93
Melidectes ochromelas 92, 100
Melidora macrorrhina 98, 105
Melilestes megarhynchus 100, 107
Meliphaga albonotata 102, 112, 114
Meliphaga analoga 94, 100, 107
Meliphaga aruensis 100
Meliphaga flavirictus 102, 112, 114
Meliphaga montana 100, 112
Meliphaga orientalis 94, 100, 112
Melipotes 113
Melipotes ater 92
Melipotes carolae 92, 100
Melipotes fumigatus 92, 113
Melipotes gymnops 92, 113
meloda, Zenaida 170
meloryphus, Euscarthmus 172
menbeki, Centropus 97, 105
menstruus, Pionus 154
meridionalis, Buteogallus 170
Merops pusillus 42
Merops superciliosus 42
mesoleuca, Conothraupis 55
mesoleuca, Elaenia 51
mesomelas. Icterus 174
mesurus, Trogon 171
metallica, Aplonis 101, 107
meyerianus, Accipiter 93, 96
meyeri, Chrysococcyx 93, 97
meyeri, Epimachus 92
meyeri, Pachycephala 89, 101
meyeri, Philemon 100
Microcerculus marginatus 156
Microdynamis parva 97
Microeca flavovirescens 99, 107
Microeca papuana 92, 99
Microphilemon fuscicapillus 248
Micropsitta bruijnii 91, 93, 97
Micropsitta keiensis 105
Micropsitta pusio 93, 97
micrura, Myrmia 171
Mimus longicaudatus 173
miniatus, Myioborus 174
Mino anais 101, 107
Mino dumontii 101, 107
minor, Fregata 118
minor, Paradisaea 101, 108
minor, Zosterops 111, 112
minutillus, Chrysococcyx 93, 97
minutus, Anous 120
Mionectes striaticollis 172
Mirafra africana 42
Mirafra angolensis 43
modesta, Psittacella 88, 91
Scientific Names Index
Bull. B.O.C. 2011 132(4)
311
molitor, Batis 43
Molothrus bonariensis 174
momota, Momotus 171
Momotus momota 171
monacha, Oenanthe 226
Monachella muelleriana 99
Monarcha axillaris 91, 99, 110
Monarcha chrysomela 99, 106
Monarcha frater 99, 106
Monarcha guttulus 99, 106
Monarcha manadensis 99, 106
Monasa morphoeus 154
Montagnii, Ortalida 223
montagnii, Penelope 223
montana, Coracina 91, 93, 98
montana, Geotrygon 165, 166
montana, Meliphaga 100, 112
montanus, Peltops 94, 101, 106
Monticola brevipes 43
monti vagus, Aeronautes 171
morio, Coracina 106
morphnoides, Hieraaetus 72, 96, 295-296
morphoeus, Monasa 154
Motacilla cinerea 98
muelleriana, Monachella 99
murina, Crateroscelis 93, 98, 106
murina, Phaeomyias 165, 167, 172
Muscicapa griseisticta 99
Muscigralla brevicauda 173
musschenbroekii, Neopsittacus 91, 97
Myadestes ralloides 157
mycteria, Jabiru 148
Myiagra alecto 99, 106
Myiarchus phaeocephalus 173
Myiarchus tuberculifer 173
Myiobius atricaudus 155
Myioborus miniatus 174
Myiodynastes bairdii 173
Myiodynastes maculatus 173
Myiopagis subplacens 172
Myiopagis viridicata 51
Myiophobus fasciatus 172
Myiotheretes striaticollis 172
Myiotriccus ornatus 140, 148
Myrmeciza 14
Myrmeciza berlepschi 4, 8, 19, 21-22
Myrmeciza exsul 7, 8, 21
Myrmeciza fortis 4-5, 10, 13, 14, 16, 17, 18, 26,
30- 35
Myrmeciza goeldii 3-6, 10, 13, 14, 16, 17, 18, 19, 26,
31- 35
Myrmeciza hyperythra 10
Myrmeciza immaculata 3-40, 13-14, 16, 18-20
Myrmeciza immaculata concepcion subsp. nov.
11
Myrmeciza laemosticta 21
Myrmeciza longipes 10
Myrmeciza melanoceps 3-6, 10, 13-14, 16-18, 19,
22, 26, 31-35
Myrmeciza nigricauda 4, 19, 21
Myrmeciza palliata 21
Myrmeciza zeledoni 3, 8, 14, 16, 18-19, 21- 23, 27,
30-35
Myrmedestes 10
Myrmelastes 10
Myrmia micrura 171
Myrmoborus 21, 22
Myrmoderus 10
Myrmophylax 10
Myrmotherula schisticolor 140, 146, 157
Myrtis fanny 171
mystacalis, Cyanocorax 173
mystacea, Hemiprocne 98, 106
Myzomela adolphinae 92-93
Myzomela cruentata 93, 100, 107
Myzomela nigrita 93, 100
Myzomela rosenbergii 90, 92-93, 100, 107
naevia, Clanga 135
naevia, Tapera 171
namaquus, Dendropicos 69
nana, Grallaricula 219
nanus, Ptilinopus 105
napensis. Megascops 188
napensis. Megascops (guatemalae) 182
napensis. Megascops [guatemalae] 187, 193
napoleonis, Polyplectron 253-254, 255-256,
257-258
Nectarinia aspasia 100
Nectarinia fusca 44
Neopsittacus musschenbroekii 91, 97
niansae, Apus 195
niger, Rhynchops 134
nigra, Melanocharis 93, 100, 107
nigrescens, Pitohui 89, 92, 100
nigricans, Sayornis 172
nigricauda, Myrmeciza 4, 19, 21
nigriceps, Saltator 174
nigriceps, Turdus 173
nigricollis, Busarellus 46, 48
nigripectus, Machaerirhynchus 91, 94, 99
nigrirostris, Macropygia 96
nigrita, Myzomela 93, 100
nigrocristatus, Basileuterus 174
nigrocyanea. Halcyon 98
nigrorufa, Crateroscelis 88, 101
nigroviridis, Tangara 157
nilotica, Gelochelidon 133, 133-135
Ninox rufa 97
Ninox theomacha 97
nipalensis, Aquila 72, 296
Nisaetus 297
nitens, Collocalia 106
niveigularis, Tyr annus 173
nobilis, Otidiphaps 96, 109
Nothocercus bonapartei 157
nouhuysi, Sericornis 89, 91, 93, 99
novaeguineae, Harpyopsis 96, 108
novaeguineae, Mearnsia 98, 102, 109, 114
novaeguineae, Toxorhamphus 100, 107
novaeguineae, Zosterops 111, 112
novaehollandiae, Accipiter 96
novaehollandiae, Anhinga 283-294
novaehollandiae, Anhinga [melanogaster] 293
novaehollandiae, Scythrops 105
Numenius tenuirostris 213
nuna, Lesbia 171
Nyctibius griseus 171
nycticorax, Nycticorax 170
Nycticorax nycticorax 170
Nyctidromus albicollis 171
Scientific Names Index
312
Bull. B.O.C. 2012 132(4)
Nystalus radiatus 154
obscurus, Lichenostomus 100
obsoletum, Camptostoma 172
obsoletus, Turdus 157
occidentals, Pezoporus 277-282, 279
occidentalis, Pezoporus (Geopsittacus) 277-282
occipitalis, Lophaetus 70
oceanicus, Oceanites 215
Oceanites oceanicus 215
ocellata, Rheinardia 258
ocellatus, Podargus 97
ochraceiventris, Leptotila 166, 170
ochromelas, Melidectes 92, 100
Ochthoeca piurae 167, 172
Ocreatus underwoodii 157
Odontophorus erythrops 144, 154
Oedistoma iliolophum 100
Oedistoma pygmaeum 100
Oenanthe finschii 226, 228, 230-231, 234
Oenanthe hispanica 230
Oenanthe leucopyga 226, 232
Oenanthe leucura 226-227, 228
Oenanthe lugens 226, 228, 230-232
Oenanthe monacha 226
Oenanthe picata 226, 232
Oenanthe pleschanka 230
Onychoprion fuscata 116, 121
opaca, Aplonis 122
Oreocharis arfaki 92, 100
orientalis, Eurystomus 106
orientalis, Meliphaga 94, 100, 112
Oriolus phaeochromus 236
Oriolus szalayi 101, 106
ornata, Thlypopsis 173
ornatus, Myiotriccus 140, 148
ornatus, Ptilinopus 87, 96, 109
ornatus, Spizaetus 140, 148
Ortalida Montagnii 223
Ortalis Columbiana 156-157
Ortalis garrula 154, 157
ortoni, Penelope 222, 222-224
Otidiphaps nobilis 96, 109
Otis tarda 213
Otus choliba 183, 185
Otus roboratus 180
Oxyruncus cristatus 152, 157
Pachycare flavogriseum 92, 94, 99
Pachycepala rufinucha 100
Pachycephala griseiceps 107
Pachycephala hyperythra 94, 100, 107, 111
Pachycephala meyeri 89, 101
Pachycephala rufinucha 92
Pachycephala schlegelii 92, 94, 100
Pachycephala simplex 100
Pachycephala soror 111
Pachycephalopsis hattamensis 99
Pachyramphus albogriseus 173
Pachyramphus polychopterus 140, 148
Pachyramphus validus 48, 52
palliata, Myrmeciza 21
pallidus, Apus 194-199, 200
palpebrosa, Gerygone 99, 106
palustris, Acrocephalus 124, 126
papa, Sarcoramphus 170
papuana, Microeca 92, 99
papuensis, Chaetorhynchus 101, 107
papuensis, Coracina 98, 106
papuensis, Eurostopodus 98
papuensis, Podargus 97, 106
papuensis, Sericornis 91, 93, 99
Parabuteo unicinctus 170
Paradigalla 93, 114
Paradisaea minor 101, 108
paradoxus, Syrrhaptes 213
Parotia berlepschi 87, 90, 92, 94, 101
Parotia carolae 92
Parotia lawesii 92
Parotia sefilata 92
Parotia wahnesi 92
Parula pitiayumi 174
Parus variegatus 263, 268
parva, Microdynamis 97
Patagioenas fasciata 170
Patagioenas speciosa 154
Patagona gigas 171
pavoninus, Dromococcyx 140, 143
pectorale, Dicaeum 100, 107
Peltops blainvillii 94, 101, 106
Peltops montanus 94, 101, 106
Penelope albipennis 162, 164
Penelope barbata 164
Penelope montagnii 223
Penelope ortoni 222, 222-224
Peneothello bimaculata 93, 99
Peneothello cryptoleuca 89, 92-93, 99
Peneothello cyanus 92-93, 99
pennatus, Hieraaetus 70, 72, 295-296
Peocilodryas hypoleuca 99
Percnostola 21
Percnostola leucostigma 8
Percnostola rufifrons 10
peregrinus, Falco 170
perlatus, Ptilinopus 96, 105
personatus, Trogon 140, 144
perspicillata, Pulsatrix 171
perspicillatus, Sericornis 89, 91, 93, 99
perstriata, Ptiloprora 92
peruanum, Glaucidium 171, 190
peruviana, Grallaricula 218
peruviana, Hypocnemis 4
peruviana, Sporophila 174
Petrochelidon rufocollaris 173
Pezoporus (Geopsittacus) occidentalis 277-282
Pezoporus occidentalis 277-282, 279
Pezoporus wallicus 277
phaeocephalus, Myiarchus 173
phaeochromus, Oriolus 236
Phaeomyias murina 165, 167, 172
Phaethon rubricauda 116, 118
Phaethomis griseogularis 171
Phaetusa simplex 134
Phalacrocorax 284
Phalacrocorax melanoleucos 105
Phalacrocorax sulcirostris 105
Pharomachrus auriceps 140, 144
Pheucticus chrysogaster 174
Pheugopedius sclateri 173
Philemon buceroides 100, 107
Philemon fuscicapilla 246
Philemon fuscicapillus 236-250, 244—245
Scientific Names Index
313
Bull. B.O.C. 2011 132(4)
Philemon meyeri 100
Philydor fuscipenne 154
Philydor rufum 140, 144, 246
Phrygilus plebejus 174
Phylloscartes superciliaris 140, 148, 151
Phylloscopus poliocephalus 91, 98, 110
picata, Oenanthe 226, 232
Picoides fumigatus 140, 144, 172
Piculus aurulentus 50
Picumnus sclateri 172
Picus canus 258
Piezorhina cinerea 174
pinon, Ducula 97, 105
Pionus menstruus 154
Pipraeidea melanonota 173
Pipra erythrocephala 155
Piranga flava 174
Piranga leucoptera 157
pitangua, Megarhynchus 52
pitiayumi, Parula 174
Pitohui 112
Pitohui cristatus 100, 107
Pitohui dichrous 94, 100, 111
Pitohui ferrugineus 100, 107
Pitohui kirhocephalus 94, 100, 111
Pitohui nigrescens 89, 92, 100
Pitta erythrogaster 98, 106
Pitta sordida 98, 106
piurae, Ochthoeca 167, 172
placens, Poecilodryas 110
placentis, Charmosyna 93, 97, 105
platensis, Embernagra 52
platyrostris, Dendrocolaptes 46, 48, 50
plebejus, Phrygilus 174
pleschanka, Oenanthe 230
plicatus, Rhyticeros 98, 106
plumbea, Polioptila 173
plumipes. Circus 221
plumipes, Falco 221
Podargus ocellatus 97
Podargus papuensis 97, 106
Poecilodryas albispecularis 88, 92, 99
Poecilodryas brachyura 99
Poecilodryas hypoleuca 107
Poecilodryas placens 110
poliocephalus, Accipiter 96, 105
poliocephalus, Phylloscopus 91, 98, 110
Polioptila plumbea 173
polychopterus, Pachyramphus 140, 148
polygrammus, Xanthotis 100
polyosoma, Buteo 170
Polyplectron bicalcaratum 251-254, 255, 257
Polyplectron chalcurum 251-253, 254, 257
Polyplectron germaini 252-253, 254-255, 257
Polyplectron inopinatum 251-252, 253-254, 257,
258
Polyplectron katsumatae 251-259, 253-254
Polyplectron malacense 251, 253, 254, 255, 256,
257-258
Polyplectron napoleonis 253-254, 255-256, 2 57-258
Polyplectron schleiermacheri 251-259, 253-254, 256
pomarina, Aquila 70-71, 135, 295
Pomatostomus isidorei 98, 107, 112
Poospiza hispaniolensis 174
porphyrocephala, Iridosornis 157
Pratincola torquata 265
Premnoplex brunnescens 157
Prinia atrogularis 258
Probosciger aterrimus 97, 105
Procellaria 211
Procellaria conspicillata 211
Progne chalybea 173
propinqua, Synallaxis 60-63, 62
Prunella fagani 204
psaltria, Carduelis 174
Psarocolius wagleri 156
Pseudelaenia leucospodia 172
Pseudeos fuscata 88, 97
Psittacella madaraszi 88, 91, 97
Psittacella modesta 88, 91
Psittaculirostris desmarestii 105
Psittaculirostris salvadorii 97
Psittrichas fulgidus 97
Pterodroma 211, 215
Pterodroma arminjoniana 211
Pterodroma incerta 211
Pterodroma macroptera 209-212, 210
Ptilinopus coronulatus 102, 109, 114
Ptilinopus iozonus 96, 105
Ptilinopus magnificus 96, 105
Ptilinopus nanus 105
Ptilinopus ornatus 87, 96, 109
Ptilinopus perlatus 96, 105
Ptilinopus pulchellus 96, 105
Ptilinopus rivoli 96, 105
Ptilinopus superbus 96, 105
Ptilinopus viridis 96
Ptiloprora 114
Ptiloprora guisei 92
Ptiloprora mayri 92, 100
Ptiloprora perstriata 92
Ptiloris magnificus 101, 108
Ptilorrhoa caerulescens 93, 98
Ptilorrhoa castanonota 93, 98, 106
Ptilorrhoa leucosticta 88, 91, 93, 98
Puffinus gravis 209
puffinus, Puffinus 215
Puffinus puffinus 215
pulchella, Charmosyna 93, 97
pulchellus, Ptilinopus 96, 105
pulcher, Melanerpes 154
Pulsatrix 178
Pulsatrix perspicillata 171
purpurata, Querula 155
pusilla, Alcedo 105
pusillus, Campylorhamphus 157
pusillus, Merops 42
pusio, Micropsitta 93, 97
Pycnonotus barbatus 78
pycnopygius, Achaetops 43
Pycnopygius ixoides 100
Pycnopygius stictocephalus 100
pygmaeum, Oedistoma 100
Pygochelidon cyanoleuca 173
Pyriglena 10, 21
Pyriglena ellisiana 7
Pyriglena leuconota 24
Pyrocephalus rubinus 172
Querula purpurata 155
quoyi, Cracticus 101, 107
Scientific Names Index
314
Bull. B.O.C. 2012 132(4)
radiatus, Nystalus 154
radjah, Tadoma 96
Rallicula forbesi 91
Rallicula leucospila 91
Rallicula mayri 91, 96
ralloides, Myadestes 157
ramobustorum subsp. nov., Eudocimus albus 128
rapax, Aquila 72, 296
reevei, Turdus 173
refectus, Buteo 221
regius, Cicinnurus 101, 107
reinwardtii, Reinwardtoena 96
reinwardt, Megapodius 105
Reinwardtoena reinwardtii 96
rex, Clytoceyx 98
Rhamphocharis crassirostris 90, 92, 100
Rheinardia ocellata 258
Rhipidura albolimbata 91, 94, 99
Rhipidura atra 91, 94, 99
Rhipidura brachyrhyncha 91, 99
Rhipidura hyperythra 94, 99, 110
Rhipidura leucothorax 99, 106
Rhipidura maculipectus 106
Rhipidura rufidorsa 99, 106
Rhipidura rufiventris 99, 106
Rhipidura threnothorax 93, 99, 106
Rhynchops niger 134
Rhynchospiza stolzmanni 174
Rhyticeros plicatus 98, 106
ridgwayi, Aegolius 178
ridgwayi, Glaucidium (brasilianum) 149
Riparia cincta 43
rivolii, Colaptes 140, 144
rivoli, Ptilinopus 96, 105
roboratus. Megascops 171, 182-184, 187-188, 193
roboratus, Otus 180
robusta, Crateroscelis 88, 91, 93, 98
roratus, Eclectus 97, 105
rosenbergii, Myzomela 90, 92-93, 100, 107
ruber, Eudocimus 128
rubicola, Saxicola 260, 261-262
Rubicola, Saxicola 267
rubiginosus, Colaptes 172
rubinus, Pyrocephalus 172
rubra, Eugerygone 92, 99
rubricauda, Phaethon 116, 118
rufa, Anhinga 283-294
rufa, Ninox 97
rufescens, Egretta 128
rufescens, Sericornis 91
ruficapilla, Grallaria 172
ruficaudus, Lanius 272
ruficollis, Chrysococcyx 88, 91, 93, 97
ruficollis, Gerygone 99, 110
ruficollis, Jynx 42
ruficollis, Stelgidopteryx 173
ruficollis, Syndactyla 166, 172
rufidorsa, Rhipidura 99, 106
rufifrons, Percnostola 10
rufigaster, Ducula 97, 105
rufigula, Gallicolumba 96
rufimarginatus, Herpsilochmus 155
rufinucha, Pachycepala 92, 100
rufiventris, Rhipidura 99, 106
rufocollaris, Petrochelidon 173
rufum, Philydor 140, 144, 146
rustica, Hirundo 173
rutila, Streptoprocne 158, 171
Saltator atripennis 15
Saltator coerulescens 156
Saltator nigriceps 174
Saltator striatipectus 174
salvadorii, Psittaculirostris 97
Salvadorina waigiuensis 87, 96
salvini, Tumbezia 172
sanctaecatarinae. Megascops 48
sanctaemariae, Cymbilaimus 66
sancta. Halcyon 105
sanguiniceps, Haematortyx 252, 254
Sarcoramphus papa 170
Sarkidiornis melanotos 164
saturata, Aethopyga 258
saturata, Scolopax 91, 96
Saxicola hemprichii 267, 268
Saxicola leucura 234
Saxicola maurus 260, 261, 262, 264, 266, 267, 268
Saxicola rubicola 260, 261-262
Saxicola Rubicola 267
Saxicola torquata 268
Saxicola torquatus 260-269
Sayornis nigricans 172
Scardafella squammata 140, 149
Schiffornis stenorhyncha 155
Schiffornis turdina 155
schisticeps, Coracina 93, 98
schisticolor, Myrmotherula 140, 146, 157
schlegelii, Pachycephala 92, 94, 100
schleiermacheri, Polyplectron 251-259, 253-254,
256
scirpaceus, Acrocephalus 124
sclateri, Pheugopedius 173
sclateri, Picumnus 172
scolopaceus, Eudynamys 97
Scolopax alba 129
Scolopax fusca 129
Scolopax saturata 91, 96
scotops, Crithagra 69
Scytalopus 4
Scytalopus atratus 140, 146, 157
Scytalopus latrans subcinereus 172
scythicus nom. nov., Charadrius leschenaultii 64
Scythrops novaehollandiae 105
seebohmi, Atlapetes 174
sefilata, Parotia 92
Seleucidis melanoleucus 101, 107
semipalmatus, Catoptrophorus 46, 48
Sericornis 89
Sericornis arfakianus 89, 91, 93, 98
Sericornis beccarii 99
Sericornis nouhuysi 89, 91, 93, 99
Sericornis papuensis 91, 93, 99
Sericornis perspicillatus 89, 91, 93, 99
Sericornis rufescens 91
Sericornis spilodera 89, 93, 98, 106
Sericornis virgatus 89, 93, 99, 110
Sericulus aureus 101
Serinus 69
Serinus capistratus 44
Serinus citrinelloides 44
serranus, Turdus 157
Scientific Names Index
Bull. B.O.C. 2011 132(4)
315
severa, Mackenziaena 66
Sicalis flaveola 174
simplex, Geoffroyus 93, 97
simplex, Pachycephala 100
simplex, Phaetusa 134
Sipia 10
Sipodotus wallacii 98
sittoides, Diglossa 173
sloetii, Campochaera 98
solitarius, Harpyhaliaetus 170
sordida. Pitta 98, 106
soror, Pachycephala 111
souleyetti, Lepidocolaptes 172
sparverius, Falco 170
speciosa, Patagioenas 154
spectabilis, Elaenia 51
speculigera, Conothraupis 55-59, 56, 173
spilodera, Sericornis 89, 93, 98, 106
spilogaster, Aquila 71-72, 296-297
spixii, Cyanopsitta 213
Spizaetus 297
Spizaetus africana 72
Spizaetus africanus 70
Spizaetus ornatus 140, 148
Sporophila 55
Sporophila peruviana 174
Sporophila telasco 174
Sporopipes squamifrons 44
squamatus, Francolinus 41
squamifrons, Sporopipes 44
squammata, Scardafella 140, 149
Steatornis caripensis 166, 171
Stelgidopteryx ruficollis 173
stenorhyncha, Schiffornis 155
stephani, Chalcophaps 96, 105
Sterna bernsteini 213
Sterna sumatrana 121
stictocephalus, Pycnopygius 100
stolidus, Anous 120
stolzmanni, Rhynchospiza 174
Streptoprocne rutila 158, 171
Streptoprocne zonaris 171
striata, Butorides 170
striaticollis, Mionectes 172
striaticollis, Myiotheretes 172
striatipectus, Saltator 174
striatus, Accpiter 170
Strix virgata 177
Sturnella belicosa 174
subalaris, Syndactyla 15, 140, 146, 152, 157
subcinereus, Scytalopus latrans 172
subcristata, Aviceda 96
subplacens, Myiopagis 172
subruficapilla, Cisticola 43
Sula 284
Sula dactylatra 116, 120
Sula leucogaster 120
sula, Sula 120
Sula sula 120
sulcirostris, Crotophaga 171
sulcirostris, Phalacrocorax 105
sulphurescens, Tolmomyias 46, 48, 52
sumatrana, Ardea 108
sumatrana. Sterna 121
superba, Lophorina 84, 92-93
superbus, Ptilinopus 96, 105
supercilaris, Drymodes 99
superciliaris, Burhinus 170
superciliaris, Cantorchilus 173
superciliaris, Phylloscartes 140, 148, 151
superciliosus, Merops 42
swainsonii, Gampsonyx 170
Sylvia crassirostris 75-83, 77, 80
Sylvia hortensis 75-83, 77, 80
Sylvia hortensis cyrenaicae, subsp. nov. 79
Synallaxis albescens 157
Synallaxis azarae 157, 172
Synallaxis gujanensis 60
Synallaxis macconnelli 60
Synallaxis propinqua 60-63, 61
Syndactyla ruficollis 166, 172
Syndactyla subalaris 15, 140, 146, 152, 157
Syrrhaptes paradoxus 213
szalayi, Oriolus 101, 106
Tachyphonus 56
Tadorna radjah 96
tahitica, Hirundo 98
Talegalla cuvieri 108
Talegalla fuscirostris 102, 104, 108, 114
Talegalla jobiensis 96
Tangara arthus 140, 148
Tangara nigroviridis 157
Tangara viridicollis 173
Tantalus coco 129
Tanysiptera galatea 98, 105
Tapera naevia 171
Taraba 66-67
Taraba major 66
tarda, Otis 213
Tchagra australis 44
telasco, Sporophila 174
telescophthalmus. Arses 106
tenebricosa, Tyto 97
teneriffae, Cyanistes 68
tenuirostris, Numenius 213
Thalasseus bergii 116,121
Thalurania 144
Thalurania colombica 140, 150, 152
Thalurania fannyi 140, 150, 152
Thalurania sp. 150, 157
Thamnophilus bernardi 172
Thamnophilus immaculatus 7
Thamnophilus unicolor 22, 157
Thamnophilus zarumae 172
theomacha, Ninox 97
Theristicus melanopis 130
Thlypopsis ornata 173
Thraupis 56
Thraupis bonariensis 173
Thraupis episcopus 173
threnothorax, Rhipidura 93, 99, 106
thula, Egretta 170
Tiaris bicolor 140, 150
Tiaris fuliginosus 140, 150, 151
Timeliopsis fulvigula 90, 92, 100
Timeliopsis griseigula 102, 112, 114
Tinamus major 153
Todirostrum cinereum 172
Tolmomyias sulphurescens 46, 48, 52
torotoro. Halcyon 98, 109
Scientific Names Index
Bull. B.O.C. 2012 132(4)
316
torquata, Megaceryle 171
torquata, Pratincola 265
torquata, Saxicola 268
torquatus, Lanioturdus 43
torquatus, Saxicola 260-269
torquatus, Turdus 79
toulsoni, Apus 42
Toxorhamphus novaeguineae 100, 107
Tregellasia leucops 99, 110
triangularis, Xiphorhynchus 157
Trichoglossus haematodus 97, 105
trichroa, Erythrura 100
trifasciatus, Basileuterus 174
tristis, Corvus 101, 107
tristissima, Lonchura 101
tristriatus, Basileuterus 20, 157
trochilirostris, Campylorhamphus 152, 157
Troglodytes aedon 173
Trogon mesurus 171
Trogon personatus 140, 144
Tropidorhynchus fuscicapillus 248
tuberculifer, Myiarchus 173
Tumbezia salvini 172
turdina, Schiffornis 155
Turdoides melanops 44
Turdus chiguanco 173
Turdus fuscater 157, 173
Turdus ignobilis 140, 148
Turdus nigriceps 173
Turdus obsoletus 157
Turdus reevei 173
Turdus serranus 157
Turdus torquatus 79
Tyrannus melancholicus 173
Tyrannus niveigularis 173
Tytoalba 171, 177
Tyto tenebricosa 97
unappendiculatus, Casuarius 96, 213
underwoodii, Ocreatus 157
unduligera, Frederickena 65
unicinctus, Parabuteo 170
unicolor, Apus 194
unicolor, Thamnophilus 22, 157
unirufus, Lipaugus 155, 157
Uropsalis lyra 140, 143
urubitinga, Buteogallus 170
validus, Pachyramphus 48, 52
vanikorensis, Collocalia 98, 106
variegatus, Parus 263, 268
variolosus, Cacomantis 97, 105
Verniliornis callonotus 172
verreauxii, Aquila 72, 296
verreauxi, Leptotila 170
versteri, Melanocharis 90, 92-93, 100
victoria, Goura 96, 213
Vireo leucophrys 140, 148
virgata, Strix 177
virgatus, Hyloclistes 140, 144, 146
virgatus, Sericornis 89, 93, 99, 110
virginianus. Bubo 177
viridicata, Myiopagis 51
viridicollis, Tangara 173
viridis, Ptilinopus 96
vociferus, Charadrius 170
Volatina jacarina 174
Vultur gryphus 170
wagleri, Aratinga 170
wagleri, Psarocolius 156
wahlbergi, Aquila 295
wahlbergi, Hieraaetus 72, 295-296
wahnesi, Parotia 92
waigiuensis, Salvadorina 87, 96
wallacii, Sipodotus 98
wallicus, Pezoporus 277
warszewiczi. Dives 174
watkinsi, Grallaria 172
weberi, Lipaugus 157
weiskei, Hieraaetus 72, 296
wilsonia, Charadrius 64
Xanthotis flavi venter 100, 107, 112
Xanthotis polygrammus 100
Xiphorhynchus erythropygius 140, 144, 146, 157
Xiphorhynchus triangularis 157
yncas, Cyanocorax 157
ypiranganus, Emberizoides 52
zarumae, Thamnophilus 172
zeledoni, Myrmeciza 3, 8, 14, 16-19, 21-23, 27,
30-33, 35
Zenaida auriculata 170
Zenaida meloda 170
zoeae, Ducula 97, 105
zonaris, Streptoprocne 171
Zonotrichia capensis 174
Zoothera dauma 98
Zosterops atrifrons 94, 100
Zosterops fuscicapilla 100
Zosterops fuscicapillus 92, 94, 111
Zosterops minor 111-112
Zosterops novaeguineae 111-112
Page 89
line 12
Page 92
lines 10
Page 93
line 36
Page 93, 98
lines 37, 31
Page 98
line 7
Page 106
line 34
Page 107
line 33
Page 157
line 37
Page 170
line 33
Page 171
line 17
Page 188
line 24
CORRECTIONS TO TEXT
Gerygone chloronota not Gerygone chloronata
Pachycephala rufinucha not Pachyephala rufinucha
Ptilorrhoa leucosticta not Ptilorrhoa leucostica
Ptilorrhoa castanonota not Ptilorrhoa castanonotus
Mearnsia novaeguineae not Mearnsia noveaguineae
Monarcha guttulus not Monarcha guttula
Seleucidis melanoleucus not Seleucidis melanoleuca
Camphylorhamphus pusillus not Camphylorhynchus pusillus
Patagioenas fasciata not Patagioena fasciata
Nyctidromus albicollis not Nyctidroma albicollis
Megascops roboratus not Megascops robotatus
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Registered Charity No. 279583
Bulletin of the British Ornithologists' Club
ISSN 0007-1595
Edited by Guy M. Kirwan
Associate Editor: Frank D. Steinheimer
Volume 132, Number 4, pages 225-316
CONTENTS
Club Announcements 225
SHIRIHAI, H. Correcting the identification of two rare wheatear records in Israel 226
BESSON, L. The collecting history and distribution of Dusky Friarbird Philemon fuscicapillus
(Morotai, Indonesia) elucidated from museum data 236
DAVISON, G. W. H., JIANG, C, ZHENGWANG, Z. & DE, C. Full tree resolution of Polyplectron
Temminck, 1813, confirms species status of Hainan P. katsumatae Rothschild, 1906, and Bornean
Peacock-Pheasants P. schleiermacheri Briiggemann, 1877 251
SVENSSON, L., SHIRIHAI, H„ FRAHNERT, S. & DICKINSON, E. C. Taxonomy and nomenclature
of the Stonechat complex Saxicola torquatus sensu lato in the Caspian region 260
PEARSON, D., SVENSSON, L. & FRAHNERT, S. Further on the type series and nomenclature of the
Isabelline Shrike Lanius isabellinus 270
BLACK, A. B. Collection localities of the Night Parrot Pezoporus ( Geopsittacus ) occidentalis
(Gould, 1861) 277
SCHODDE, R., KIRWAN, G. M. & PORTER, R. Morphological differentiation and speciation among
darters (Anhinga) 283
CLARK, W. S. The eagle genus Hieraaetus is distinct from Aquila, with comments on the name
Ayres' Eagle 295
Index for Volume 132 (2012) 299
EDITORIAL BOARD
Murray Bruce, R, T. Chesser, Edward C. Dickinson, Frangoise Dowsett-Lemaire, Steven M. S. Gregory, Jose
Fernando Pacheco, Robert B. Payne, Pamela C. Rasmussen, Cees Roselaar, Thomas S. Schulenberg, Lars
Svensson
Authors are invited to submit papers on topics relating to the broad themes of taxonomy, nomenclature and
distribution of birds. Descriptions of new species are especially welcome and will be given priority to ensure
rapid publication, subject to successful passage through the normal peer review procedure, and wherever
possible should be accompanied by colour photographs or paintings. On submission, manuscripts, double-
spaced and with wide margins, should be sent to the Editor, Guy Kirwan, by e-mail, to GMKirwan@aol.com.
Alternatively, two copies of manuscripts, typed on one side of the paper, may be submitted to the Editor, 74
Waddington Street, Norwich NR2 4JS, UK. Where appropriate half-tone photographs may be included and,
where necessary to illustrate important points, the Editor will consider the inclusion of colour figures. Review,
return of manuscripts for revision and subsequent stages of the publication process will be undertaken
electronically.
For instructions on style, see the inside rear cover of Bulletin 132(1) or the BOC website.
Registered Charity No. 279583
www.boc-online.org
Printed on acid-free paper.
Published by the British Ornithologists' Club
Typeset by Alcedo Publishing of Arizona, USA, and printed by Latimer Trend, UK
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